Biology, 9th Edition

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Biology NINTH EDITION

ELDRA P. SOLOMON former affiliations: University of South Florida, Tampa Hillsborough Community College

LINDA R. BERG former affiliations: University of Maryland, College Park St. Petersburg College

DIANA W. MARTIN Rutgers University

Australia • Brazil • Japan • Korea • Mexico • Singapore • Spain • United Kingdom • United States

Biology, Ninth Edition Eldra P. Solomon, Linda R. Berg, and Diana W. Martin Publisher, Life Sciences: Yolanda Cossio

© 2011, 2008 Brooks/Cole, Cengage Learning

Developmental Editor: Christopher Delgado Assistant Editor: Alexis Glubka Editorial Assistant: Joshua W. Taylor Media Editor: Lauren Oliveira Marketing Manager: Tom Ziolkowski Marketing Assistant: Elizabeth Wong Marketing Communications Manager: Linda Yip

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Student Edition: ISBN-13: 978-0-538-74125-5 ISBN-10: 0-538-74125-2

Cover Designer: John Walker Cover Image: Leafy sea dragon (Phycodurus eques) in Southern Ocean; Michael Aw/Getty Images The leafy sea dragon inhabits kelp-covered rocky reefs and seaweed beds in the waters off the southern and western coast of Australia. This fish lives in depths from about 3 m to 50 m. It feeds on plankton, mainly tiny crustaceans, and typically grows to a length of about 35 centimeters (14 inches). The leafy sea dragon, with its long leaflike extensions, blends with surrounding seaweed, a striking example of camouflage. When the male leafy sea dragon is ready to mate, his tail becomes bright yellow. Females then deposit up to 250 bright pink eggs in a spongy brood patch on the underside of the male’s tail. The eggs are fertilized during the transfer from the female to the male. The male incubates the eggs for four to six weeks. Then, miniature leafy sea dragons are released into the water. Leafy sea dragon populations have decreased due to habitat loss, pollution, and collectors who sell them for aquariums. They are currently listed as “near threatened” in the International Union for Conservation of Nature’s status list.

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Loose-leaf Edition: ISBN-13: 978-0-538-49419-9 ISBN-10: 0-538-49419-0 Brooks/Cole 20 Davis Drive Belmont, CA 94002-3098 USA Cengage Learning is a leading provider of customized learning solutions with office locations around the globe, including Singapore, the United Kingdom, Australia, Mexico, Brazil and Japan. Locate your local office at www.cengage.com/global Cengage Learning products are represented in Canada by Nelson Education, Ltd. To learn more about Brooks/Cole, visit www.cengage.com/ brookscole Purchase any of our products at your local college store or at our preferred online store www.cengagebrain.com.

DEDICATION To our families, friends, and colleagues who gave freely of their love, support, knowledge, and time as we prepared this ninth edition of Biology. Especially to Freda, Kathleen, Mical, Amy, and Belicia Alan and Jennifer Chuck and Margaret

ABOUT THE AUTHORS

ELDRA P. SOLOMON has written several leading college-level textbooks in biology and in human anatomy and physiology. Her books have been translated into more than 10 languages. Dr. Solomon earned an M.S. from the University of Florida and an M.A. and Ph.D. from the University of South Florida. Dr. Solomon taught biology and nursing students for more than 20 years. In addition to being a biologist and science author, Dr. Solomon is a biopsychologist with a special interest in the neuro-physiology of traumatic experience. Her research has focused on the neurological, endocrine, and psychological effects of trauma, including Post-traumatic Stress Disorder and development of maladaptive coping strategies. Dr. Solomon has presented her research at many national and international conferences, and her work has been published in leading professional journals. Dr. Solomon has been profiled more than 30 times in leading publications, including Who’s Who in America, Who’s Who in Science and Engineering, Who’s Who in Medicine and Healthcare, Who’s Who in American Education, Who’s Who of American Women, and Who’s Who in the World.

LINDA R. BERG is an award-winning teacher and textbook author. She received a B.S. in science education, an M.S. in botany, and a Ph.D. in plant physiology from the University of Maryland. Her research focused on the evolutionary implications of steroid biosynthetic pathways in various organisms. Dr. Berg taught at the University of Maryland at College Park for 17 years and at St. Petersburg College in Florida for 8 years. During her career, she taught introductory courses in biology, botany, and environmental science to thousands of students. At the University of Maryland, she received numerous teaching and service awards. Dr. Berg is also the recipient of many national and regional awards, including the National Science Teachers Association Award for Innovations in College Science Teaching, the Nation’s Capital Area Disabled Student Services Award, and the Washington Academy of Sciences Award in University Science Teaching. During her career as a professional science writer, Dr. Berg has authored or co-authored several leading college science textbooks. Her writing reflects her teaching style and love of science.

DIANA W. MARTIN is the Director of General Biology, Division of Life Sciences, at Rutgers University, New Brunswick Campus. She received an M.S. at Florida State University, where she studied the chromosomes of related plant species to understand their evolutionary relationships. She earned a Ph.D. at the University of Texas at Austin, where she studied the genetics of the fruit fly, Drosophila melanogaster, and then conducted postdoctoral research at Princeton University. She has taught General Biology and other courses at Rutgers for more than 25 years and has been involved in writing textbooks since 1988. She is immensely grateful that her decision to study biology in college has led to a career that allows her many ways to share her excitement about all aspects of biology.

Brief Contents

Preface

xxv

To the Student

xxxi

Part 1 THE ORGANIZATION OF LIFE 1

A View of Life

2

Atoms and Molecules: The Chemical Basis of Life

1

1

307

15

DNA Technology and Genomics

16

Human Genetics and the Human Genome

17

Developmental Genetics

323 347

369

26

The Chemistry of Life: Organic Compounds

4

Organization of the Cell

5

Biological Membranes

6

Cell Communication

46

18

Introduction to Darwinian Evolution

391

74

19

Evolutionary Change in Populations

411

106

20

Speciation and Macroevolution

21

The Origin and Evolutionary History of Life

22

The Evolution of Primates

134

Part 2 ENERGY TRANSFER THROUGH LIVING SYSTEMS 154 7

Energy and Metabolism

8

How Cells Make ATP: Energy-Releasing Pathways

172 193

Part 3 THE CONTINUITY OF LIFE: GENETICS 213 10

Chromosomes, Mitosis, and Meiosis

11

The Basic Principles of Heredity

12

DNA: The Carrier of Genetic Information

13

Gene Expression

426

465

Part 5 THE DIVERSITY OF LIFE

154

Photosynthesis: Capturing Light Energy

282

Gene Regulation

Part 4 THE CONTINUITY OF LIFE: EVOLUTION 391

3

9

14

213

237

481

23

Understanding Diversity: Systematics

24

Viruses and Subviral Agents

25

Bacteria and Archaea

26

Protists

27

Seedless Plants

28

Seed Plants

29

The Fungi

30

An Introduction to Animal Diversity

31

Sponges, Cnidarians, Ctenophores, and Protostomes 640

32

The Deuterostomes

263

481

501

517

537 561

582 601

675

626

446

Part 6 STRUCTURE AND LIFE PROCESSES IN PLANTS 708 33

Plant Structure, Growth, and Development

34

Leaf Structure and Function

35

Stem Structure and Transport

36

Roots and Mineral Nutrition

37

Reproduction in Flowering Plants

38

Plant Developmental Responses to External and Internal Signals 803

49

Endocrine Regulation

50

Reproduction

51

Animal Development

52

Animal Behavior

1052

1077 1106

708 1127

728 744

Part 8 THE INTERACTIONS OF LIFE: ECOLOGY 1153

761 781

Part 7 STRUCTURE AND LIFE PROCESSES IN ANIMALS 821 39

Animal Structure and Function: An Introduction

40

Protection, Support, and Movement

41

Neural Signaling

42

Neural Regulation

43

Sensory Systems

44

Internal Transport

45

The Immune System: Internal Defense

46

Gas Exchange

47

Processing Food and Nutrition

48

Osmoregulation and Disposal of Metabolic Wastes 1034

53

Introduction to Ecology: Population Ecology

54

Community Ecology

55

Ecosystems and the Biosphere

56

Ecology and the Geography of Life

57

Biological Diversity and Conservation Biology

1173 1196 1218

821 Appendix A

Periodic Table of the Elements

A-1

Appendix B

Classification of Organisms

Appendix C

Understanding Biological Terms

Appendix D

Abbreviations

Appendix E

Answers to Test Your Understanding Questions A-11

842 A-2

860 A-6

882 A-9

911 937

993

963

Glossary Index

1012

I-1

G-1

1153

1242

Table of Contents

Part 1 THE ORGANIZATION OF LIFE 1

A View of Life

1

Three Basic Themes

2

Characteristics of Life 2 Organisms are composed of cells 2 Organisms grow and develop 3 Organisms regulate their metabolic processes 3 Organisms respond to stimuli 4 Organisms reproduce 4 Populations evolve and become adapted to the environment 5 Levels of Biological Organization 5 Organisms have several levels of organization 6 Several levels of ecological organization can be identified 6 Information Transfer 6 DNA transmits information from one generation to the next 6 Information is transmitted by chemical and electrical signals 8 The Energy of Life

A theory is supported by tested hypotheses 19 Many hypotheses cannot be tested by direct experiment 19 Paradigm shifts allow new discoveries 21 Systems biology integrates different levels of information 21 Science has ethical dimensions 22

1

8

Evolution: The Basic Unifying Concept of Biology 10 Biologists use a binomial system for naming organisms 11 Taxonomic classification is hierarchical 11 The tree of life includes three domains and several kingdoms 11 Species adapt in response to changes in their environment 14 Natural selection is an important mechanism by which evolution proceeds 14 Populations evolve as a result of selective pressures from changes in their environment 15 The Process of Science 15 Science requires systematic thought processes 16 Scientists make careful observations and ask critical questions 16 Chance often plays a role in scientific discovery 17 A hypothesis is a testable statement 17 Many predictions can be tested by experiment 18 Researchers must avoid bias 19 Scientists interpret the results of experiments and make conclusions 19

2

Atoms and Molecules: The Chemical Basis of Life

26

Elements and Atoms 27 An atom is uniquely identified by its number of protons 27 Protons plus neutrons determine atomic mass 28 Isotopes of an element differ in number of neutrons 29 Electrons move in orbitals corresponding to energy levels 30 Chemical Reactions 31 Atoms form compounds and molecules 31 Simplest, molecular, and structural chemical formulas give different information 31 One mole of any substance contains the same number of units 31 Chemical equations describe chemical reactions 32 Chemical Bonds 32 In covalent bonds electrons are shared 32 Ionic bonds form between cations and anions 34 Hydrogen bonds are weak attractions 36 van der Waals interactions are weak forces 36 Redox Reactions

36

Water 37 Hydrogen bonds form between water molecules 37 Water molecules interact with hydrophilic substances by hydrogen bonding 38 Water helps maintain a stable temperature 38 Acids, Bases, and Salts 40 pH is a convenient measure of acidity 41 Buffers minimize pH change 42 An acid and a base react to form a salt 42

3

The Cell Nucleus

The Chemistry of Life: Organic Compounds 46

Organelles in the Cytoplasm 89 Ribosomes manufacture proteins 89 The endoplasmic reticulum is a network of internal membranes 89 The Golgi complex processes, sorts, and modifies proteins 91 Lysosomes are compartments for digestion 93 Vacuoles are large, fluid-filled sacs with a variety of functions 93 Peroxisomes metabolize small organic compounds 93 Mitochondria and chloroplasts are energy-converting organelles 94 Mitochondria make ATP through cellular respiration 94 Chloroplasts convert light energy to chemical energy through photosynthesis 96

Carbon Atoms and Organic Molecules 47 Isomers have the same molecular formula but different structures 48 Functional groups change the properties of organic molecules 48 Many biological molecules are polymers 51 Carbohydrates 51 Monosaccharides are simple sugars 52 Disaccharides consist of two monosaccharide units 53 Polysaccharides can store energy or provide structure 53 Some modified and complex carbohydrates have special roles 55 Lipids 56 Triacylglycerol is formed from glycerol and three fatty acids 57 Saturated and unsaturated fatty acids differ in physical properties 57 Phospholipids are components of cell membranes 58 Carotenoids and many other pigments are derived from isoprene units 59 Steroids contain four rings of carbon atoms 59 Some chemical mediators are lipids 59 Proteins 60 Amino acids are the subunits of proteins 60 Peptide bonds join amino acids 61 Proteins have four levels of organization 61 The amino acid sequence of a protein determines its conformation 66

4

86

The Cytoskeleton 97 Microtubules are hollow cylinders 97 Centrosomes and centrioles function in cell division 98 Cilia and flagella are composed of microtubules 98 Microfilaments consist of intertwined strings of actin 99 Intermediate filaments help stabilize cell shape 101 Cell Coverings

5

101

Biological Membranes

106

Nucleic Acids 68 Some nucleotides are important in energy transfers and other cell functions 68

The Structure of Biological Membranes 107 Phospholipids form bilayers in water 107 The fluid mosaic model explains membrane structure 108 Biological membranes are two-dimensional fluids 109 Biological membranes fuse and form closed vesicles 111 Membrane proteins include integral and peripheral proteins 111 Proteins are oriented asymmetrically across the bilayer 113

Identifying Biological Molecules

Overview of Membrane Protein Functions

Organization of the Cell

69

74

114

Cell Membrane Structure and Permeability 115 Biological membranes present a barrier to polar molecules 115 Transport proteins transfer molecules across membranes 115

The Cell: Basic Unit of Life 75 The cell theory is a unifying concept in biology 75 The organization of all cells is basically similar 75 Cell size is limited 75 Cell size and shape are adapted to function 76

Passive Transport 116 Diffusion occurs down a concentration gradient 116 Osmosis is diffusion of water across a selectively permeable membrane 117 Facilitated diffusion occurs down a concentration gradient 119

Methods for Studying Cells 77 Light microscopes are used to study stained or living cells 77 Electron microscopes provide a high-resolution image that can be greatly magnified 79 Biologists use biochemical techniques to study cell components 79

Active Transport 121 Active transport systems “pump” substances against their concentration gradients 121 Carrier proteins can transport one or two solutes 123 Cotransport systems indirectly provide energy for active transport 123

Prokaryotic and Eukaryotic Cells 81 Organelles of prokaryotic cells are not surrounded by membranes 81 Membranes divide the eukaryotic cell into compartments 82

Exocytosis and Endocytosis 123 In exocytosis, vesicles export large molecules 123 In endocytosis, the cell imports materials 125

Cell Junctions 127 Anchoring junctions connect cells of an epithelial sheet 127 Tight junctions seal off intercellular spaces between some animal cells 128 Gap junctions allow the transfer of small molecules and ions 129 Plasmodesmata allow certain molecules and ions to move between plant cells 129

6

Cell Communication

134

Cell Communication: An Overview Sending Signals

Energy and Metabolism 156 Enthalpy is the total potential energy of a system 157 Free energy is available to do cell work 157 Chemical reactions involve changes in free energy 157 Free energy decreases during an exergonic reaction 157 Free energy increases during an endergonic reaction 157 Diffusion is an exergonic process 158 Free-energy changes depend on the concentrations of reactants and products 158 Cells drive endergonic reactions by coupling them to exergonic reactions 158

135

ATP, the Energy Currency of the Cell 159 ATP donates energy through the transfer of a phosphate group 160 ATP links exergonic and endergonic reactions 160 The cell maintains a very high ratio of ATP to ADP 160

136

Reception 137 Cells regulate reception 138 Three types of receptors occur on the cell surface 139 Some receptors are located inside the cell 141

Energy Transfer in Redox Reactions 161 Most electron carriers transfer hydrogen atoms 161

Signal Transduction 141 Signaling molecules can act as molecular switches 141 Ion channel–linked receptors open or close channels 141 G protein–linked receptors initiate signal transduction 142 Second messengers are intracellular signaling agents 143 Many enzyme-linked receptors activate protein kinase signaling pathways 146 Many activated intracellular receptors are transcription factors 147 Scaffold proteins increase efficiency 147 Signals can be transmitted in more than one direction 147 Responses to Signals 147 Ras pathways involve tyrosine kinase receptors and G proteins 148 The response to a signal is amplified 148 Signals must be terminated 149 Evolution of Cell Communication

150

Enzymes 162 All reactions have a required energy of activation 162 An enzyme lowers a reaction’s activation energy 163 An enzyme works by forming an enzyme–substrate complex 163 Enzymes are specific 164 Many enzymes require cofactors 164 Enzymes are most effective at optimal conditions 165 Enzymes are organized into teams in metabolic pathways 166 The cell regulates enzymatic activity 166 Enzymes are inhibited by certain chemical agents 168 Some drugs are enzyme inhibitors 168

8

How Cells Make ATP: Energy-Releasing Pathways 172 Redox Reactions

Part 2 ENERGY TRANSFER THROUGH LIVING SYSTEMS 154 7

Energy and Metabolism

154

Biological Work 155 Organisms carry out conversions between potential energy and kinetic energy 155 The Laws of Thermodynamics 155 The total energy in the universe does not change 155 The entropy of the universe is increasing 156

173

The Four Stages of Aerobic Respiration 173 In glycolysis, glucose yields two pyruvates 175 Pyruvate is converted to acetyl CoA 177 The citric acid cycle oxidizes acetyl CoA 177 The electron transport chain is coupled to ATP synthesis 177 Inquiring About Electron Transport and Heat Aerobic respiration of one glucose yields a maximum of 36 to 38 ATPs 184 Cells regulate aerobic respiration 186 Energy Yield of Nutrients Other Than Glucose Anaerobic Respiration and Fermentation Alcohol fermentation and lactate fermentation are inefficient 188

187

183

186

9

Photosynthesis: Capturing Light Energy Light and Photosynthesis

Regulation of the Cell Cycle

193

194

Chloroplasts 195 Chlorophyll is found in the thylakoid membrane 195 Chlorophyll is the main photosynthetic pigment 196 Overview of Photosynthesis 198 ATP and NADPH are the products of the light-dependent reactions: An overview 198 Carbohydrates are produced during the carbon fixation reactions: An overview 199

Sexual Life Cycles

The Light-Dependent Reactions 199 Photosystems I and II each consist of a reaction center and multiple antenna complexes 200 Noncyclic electron transport produces ATP and NADPH 200 Cyclic electron transport produces ATP but no NADPH 202 ATP synthesis occurs by chemiosmosis 202

11

208

Photosynthesis in Plants and in the Environment 209

Chromosomes, Mitosis, and Meiosis

The Basic Principles of Heredity

237

Using Probability to Predict Mendelian Inheritance 247

Part 3 THE CONTINUITY OF LIFE: GENETICS 213 10

233

Mendel’s Principles of Inheritance 238 Alleles separate before gametes are formed: the principle of segregation 241 Alleles occupy corresponding loci on homologous chromosomes 242 A monohybrid cross involves individuals with different alleles of a given locus 242 A dihybrid cross involves individuals that have different alleles at two loci 245 Alleles on nonhomologous chromosomes are randomly distributed into gametes: the principle of independent assortment 245 Recognition of Mendel’s work came during the early 20th century 246

The Carbon Fixation Reactions 204 Most plants use the Calvin cycle to fix carbon 204 Photorespiration reduces photosynthetic efficiency 206 The initial carbon fixation step differs in C4 plants and in CAM plants 206 CAM plants fix CO2 at night 207 Metabolic Diversity

223

Sexual Reproduction and Meiosis 225 Meiosis produces haploid cells with unique gene combinations 227 Prophase I includes synapsis and crossing-over 230 During meiosis I, homologous chromosomes separate 231 Chromatids separate in meiosis II 231 Mitosis and meiosis lead to contrasting outcomes 231

Inquiring About Solving Genetic Problems The rules of probability can be applied to a variety of calculations 248

Inheritance and Chromosomes 249 Linked genes do not assort independently 249 Calculating the frequency of crossing-over reveals the linear order of linked genes on a chromosome 250 Sex is generally determined by sex chromosomes 251

213

Eukaryotic Chromosomes 214 DNA is organized into informational units called genes 214 DNA is packaged in a highly organized way in chromosomes 214 Chromosome number and informational content differ among species 215 The Cell Cycle and Mitosis 217 Chromosomes duplicate during interphase 217 During prophase, duplicated chromosomes become visible with the microscope 218 Prometaphase begins when the nuclear envelope breaks down 220 Duplicated chromosomes line up on the midplane during metaphase 220 During anaphase, chromosomes move toward the poles 221 During telophase, two separate nuclei form 221 Cytokinesis forms two separate daughter cells 222 Mitosis produces two cells genetically identical to the parent cell 222 Lacking nuclei, prokaryotes divide by binary fission 222

248

Extensions of Mendelian Genetics 255 Dominance is not always complete 255 Multiple alleles for a locus may exist in a population 256 A single gene may affect multiple aspects of the phenotype 257 Alleles of different loci may interact to produce a phenotype 257 In polygenic inheritance, the offspring exhibit a continuous variation in phenotypes 257 Genes interact with the environment to shape phenotype 258

12

DNA: The Carrier of Genetic Information

263

Evidence of DNA as the Hereditary Material DNA is the transforming principle in bacteria 264 DNA is the genetic material in certain viruses 265

264

The Structure of DNA 265 Nucleotides can be covalently linked in any order to form long polymers 267 DNA is made of two polynucleotide chains intertwined to form a double helix 268 In double-stranded DNA, hydrogen bonds form between A and T and between G and C 269 DNA Replication 271 Meselson and Stahl verified the mechanism of semiconservative replication 271 Semiconservative replication explains the perpetuation of mutations 273 DNA replication requires protein “machinery” 273 Enzymes proofread and repair errors in DNA 276 Telomeres cap eukaryotic chromosome ends 277

13

Gene Expression

Mutations 300 Base-pair substitution mutations result from the replacement of one base pair by another 301 Frameshift mutations result from the insertion or deletion of base pairs 301 Some mutations involve mobile genetic elements 303 Mutations have various causes 303

14

307

Gene Regulation in Bacteria and Eukaryotes: An Overview 308 Gene Regulation in Bacteria 309 Operons in bacteria facilitate the coordinated control of functionally related genes 309 Some posttranscriptional regulation occurs in bacteria 313 Gene Regulation in Eukaryotic Cells 314 Eukaryotic transcription is controlled at many sites and by many regulatory molecules 315 The mRNAs of eukaryotes have many types of posttranscriptional control 319 Posttranslational chemical modifications may alter the activity of eukaryotic proteins 320

282

Discovery of the Gene–Protein Relationship Beadle and Tatum proposed the one-gene, one-enzyme hypothesis 283

Gene Regulation

283

Information Flow from DNA to Protein: An Overview 285 DNA is transcribed to form RNA 285 RNA is translated to form a polypeptide 285 Biologists cracked the genetic code in the 1960s 287 The genetic code is virtually universal 288 The genetic code is redundant 288 Transcription 288 The synthesis of mRNA includes initiation, elongation, and termination 289 Messenger RNA contains base sequences that do not directly code for protein 290 Eukaryotic mRNA is modified after transcription and before translation 291 Translation 293 An amino acid is attached to tRNA before incorporation into a polypeptide 293 The components of the translational machinery come together at the ribosomes 294 Translation begins with the formation of an initiation complex 294 During elongation, amino acids are added to the growing polypeptide chain 295 One of three stop codons signals the termination of translation 296 Variations in Gene Expression 297 Transcription and translation are coupled in bacteria 297 Biologists debate the evolution of eukaryotic gene structure 298 Several kinds of eukaryotic RNA have a role in gene expression 298 The definition of a gene has evolved as biologists have learned more about genes 300 The usual direction of information flow has exceptions 300

15

DNA Technology and Genomics

323

DNA Cloning 324 Restriction enzymes are “molecular scissors” 324 Recombinant DNA forms when DNA is spliced into a vector 324 DNA can be cloned inside cells 326 A cDNA library is complementary to mRNA and does not contain introns 329 The polymerase chain reaction amplifies DNA in vitro 329 DNA Analysis 331 Gel electrophoresis is used for separating macromolecules 331 DNA, RNA, and protein blots detect specific fragments 331 Restriction fragment length polymorphisms are a measure of genetic relationships 331 Methods to rapidly sequence DNA have been developed 333 Genomics 335 Identifying protein-coding genes is useful for research and for medical applications 336 One way to study gene function is to silence genes one at a time 336 DNA microarrays are a powerful tool for studying gene expression 336 The Human Genome Project stimulated studies of genome sequencing for other species 338 Several scientific fields—bioinformatics, pharmacogenetics, and proteomics—have emerged 338

A totipotent nucleus contains all the instructions for development 372 The first cloned mammal was a sheep 373 Stem cells divide and give rise to differentiated cells 373

Applications of DNA Technologies 339 DNA technology has revolutionized medicine 339 DNA fingerprinting has numerous applications 340 Transgenic organisms have foreign DNA incorporated into their cells 340 DNA Technology Has Raised Safety Concerns

16

The Genetic Control of Development 375 A variety of model organisms provide insights into basic biological processes 376 Many genes that control development have been identified in the fruit fly 376 Caenorhabditis elegans has a relatively rigid developmental pattern 381 The mouse is a model for mammalian development 384 Arabidopsis is a model for studying plant development, including transcription factors 386

343

Human Genetics and the Human Genome 347 Studying Human Genetics 348 Human chromosomes are studied by karyotyping 348 Family pedigrees help identify certain inherited conditions 349 The Human Genome Project sequenced the DNA on all human chromosomes 349 Comparative genomics has revealed DNA identical in both mouse and human genomes 351 Researchers use mouse models to study human genetic diseases 351 Abnormalities in Chromosome Number and Structure 351 Down syndrome is usually caused by trisomy 21 352 Most sex chromosome aneuploidies are less severe than autosomal aneuploidies 354 Abnormalities in chromosome structure cause certain disorders 355 Genomic imprinting is determined by whether inheritance is from the male or female parent 356

Cancer and Cell Development

Part 4 THE CONTINUITY OF LIFE: EVOLUTION 391 18

17

Developmental Genetics

369

Cell Differentiation and Nuclear Equivalence 370 Most cell differences are due to differential gene expression 370

391

392

Pre-Darwinian Ideas about Evolution

392

Darwin and Evolution 393 Darwin proposed that evolution occurs by natural selection 395 The modern synthesis combines Darwin’s theory with genetics 395 Biologists study the effect of chance on evolution 396 Evidence for Evolution 397 The fossil record provides strong evidence for evolution 397 The distribution of plants and animals supports evolution 400 Comparative anatomy of related species demonstrates similarities in their structures 402 Molecular comparisons among organisms provide evidence for evolution 404 Developmental biology helps unravel evolutionary patterns 406 Evolutionary hypotheses are tested experimentally 407

Gene Therapy 361 Gene therapy programs are carefully scrutinized 361

Human Genetics, Society, and Ethics 365 Genetic discrimination provokes heated debate 365 Many ethical issues related to human genetics must be addressed 366

Introduction to Darwinian Evolution What Is Evolution?

Genetic Diseases Caused by Single-Gene Mutations 358 Many genetic diseases are inherited as autosomal recessive traits 358 Some genetic diseases are inherited as autosomal dominant traits 360 Some genetic diseases are inherited as X-linked recessive traits 361

Genetic Testing and Counseling 362 Prenatal diagnosis detects chromosome abnormalities and gene defects 362 Genetic screening searches for genotypes or karyotypes 364 Genetic counselors educate people about genetic diseases 364

387

19

Evolutionary Change in Populations

411

Genotype, Phenotype, and Allele Frequencies

412

The Hardy–Weinberg Principle 412 Genetic equilibrium occurs if certain conditions are met 414 Human MN blood groups are a valuable illustration of the Hardy–Weinberg principle 414 Microevolution 415 Nonrandom mating changes genotype frequencies 415 Mutation increases variation within a population 416 In genetic drift, random events change allele frequencies 416 Gene flow generally increases variation within a population 417 Natural selection changes allele frequencies in a way that increases adaptation 417

Genetic Variation in Populations 420 Genetic polymorphism can be studied in several ways 420 Balanced polymorphism exists for long periods 420 Neutral variation may give no selective advantage or disadvantage 422 Populations in different geographic areas often exhibit genetic adaptations to local environments 422

20

Speciation and Macroevolution

426

What Is a Species? 427 The biological species concept is based on reproductive isolation 427 The phylogenetic species concept defines species based on such evidence as molecular sequencing 427

The History of Life 455 Rocks from the Ediacaran period contain fossils of cells and simple animals 455 A diversity of organisms evolved during the Paleozoic era 455 Dinosaurs and other reptiles dominated the Mesozoic era 458 Inquiring About The Origin of Flight in Birds The Cenozoic era is the age of mammals 461

22

Primate Adaptations

21

Inquiring About The Smallest Humans 476 Scientists have reached a near consensus on the origin of modern Homo sapiens 476

438

The Origin and Evolutionary History of Life 446 Chemical Evolution on Early Earth 447 Organic molecules formed on primitive Earth 448 The First Cells 449 The origin of a simple metabolism within a membrane boundary may have occurred early in the evolution of cells 449 Molecular reproduction was a crucial step in the origin of cells 450 Biological evolution began with the first cells 451 The first cells were probably heterotrophic 452 Aerobes appeared after oxygen increased in the atmosphere 453 Eukaryotic cells descended from prokaryotic cells 453

466

Hominin Evolution 471 The earliest hominins may have lived 6 mya to 7 mya 472 Ardipithecus, Australopithecus, and Paranthropus are australopithecines, or “southern man apes” 472 Homo habilis is the oldest member of genus Homo 474 Homo ergaster may have arisen from H. habilis 474 Homo erectus probably evolved from H. ergaster 474 Archaic humans date from about 1.2 mya to 200,000 years ago 475 Neandertals appeared approximately 250,000 years ago 475

Speciation 430 Long physical isolation and different selective pressures result in allopatric speciation 432 Two populations diverge in the same physical location by sympatric speciation 434 The study of hybrid zones has made important contributions to what is known about speciation 437

Macroevolution 439 Evolutionary novelties originate through modifications of pre-existing structures 440 Adaptive radiation is the diversification of an ancestral species into many species 440 Extinction is an important aspect of evolution 441 Is microevolution related to speciation and macroevolution? 442

465

Primate Classification 467 Suborder Anthropoidea includes monkeys, apes, and humans 467 Apes are our closest living relatives 469

Reproductive Isolation 428 Prezygotic barriers interfere with fertilization 428 Postzygotic barriers prevent gene flow when fertilization occurs 430 Biologists are discovering genes responsible for reproductive isolating mechanisms 430

The Rate of Evolutionary Change

The Evolution of Primates

461

Cultural Change 478 Development of agriculture resulted in a more dependable food supply 478 Human culture has had a profound impact on the biosphere 478

Part 5 THE DIVERSITY OF LIFE 23

481

Understanding Diversity: Systematics

481

Classifying Organisms 482 Organisms are named using a binomial system 482 Each taxonomic level is more general than the one below it 483 Determining the Major Branches in the Tree of Life 483 Systematics is an evolving science 483 The three domains form the three main branches of the tree of life 485 Some biologists are moving away from Linnaean categories 486 Phylogenetic trees show hypothesized evolutionary relationships 486 Systematists continue to consider other hypotheses 487

Reconstructing Evolutionary History 488 Homologous structures are important in determining evolutionary relationships 489 Shared derived characters provide clues about phylogeny 489 Biologists carefully choose taxonomic criteria 489 Molecular homologies help clarify phylogeny 490 Taxa are grouped based on their evolutionary relationships 491

Prokaryote Reproduction and Evolution 521 Rapid reproduction contributes to prokaryote success 522 Prokaryotes transfer genetic information 522 Evolution proceeds rapidly in bacterial populations 523 Nutritional and Metabolic Adaptations 524 Most prokaryotes require oxygen 524 Some prokaryotes fix and metabolize nitrogen 525

Constructing Phylogenetic Trees 493 Outgroup analysis is used in constructing and interpreting cladograms 493 A cladogram is constructed by considering shared derived characters 494 In a cladogram each branch point represents a major evolutionary step 494 Systematists use the principles of parsimony and maximum likelihood to make decisions 496

24

Viruses and Subviral Agents

The Phylogeny of the Two Prokaryote Domains 525 Key characters distinguish the three domains 525 Taxonomy of archaea and bacteria continuously changes 526 Many archaea inhabit harsh environments 526 Bacteria are the most familiar prokaryotes 527 Impact on Ecology, Technology, and Commerce Prokaryotes form intimate relationships with other organisms 530 Prokaryotes play key ecological roles 530 Prokaryotes are important in many commercial processes and in technology 531

501

The Status and Structure of Viruses 502 Viruses are very small 502 A virus consists of nucleic acid surrounded by a protein coat 502 The capsid is a protective protein coat 502 Some viruses are surrounded by an envelope 503 Classification of Viruses

Viral Diseases 506 Viruses cause serious plant diseases 506 Viruses cause serious diseases in animals 507 Inquiring About Influenza and Other Emerging Diseases 509 512

Subviral Agents 513 Satellites depend on helper viruses 513 Viroids are the smallest known pathogens 513 Prions are protein particles 513

25

Bacteria and Disease 532 Many scientists have contributed to our understanding of infectious disease 532 Many adaptations contribute to pathogen success 532 Antibiotic resistance is a major public health problem 534

504

Viral Replication 504 Bacteriophages infect bacteria 505 Viruses replicate inside host cells 505

Evolution of Viruses

530

Bacteria and Archaea

517

The Structure of Bacteria and Archaea 518 Prokaryotes have several common shapes 518 Prokaryotic cells do not have membrane-enclosed organelles 518 A cell wall protects most prokaryotes 519 Some bacteria produce capsules or slime layers 519 Some prokaryotes have fimbriae or pili 519 Some bacteria survive unfavorable conditions by forming endospores 520 Many types of prokaryotes are motile 520

26

Protists

537

Diversity in the Protists

538

How Did Eukaryotes Evolve? 539 Mitochondria and chloroplasts probably originated from endosymbionts 539 A consensus in eukaryote classification is beginning to emerge 539 Excavates 540 Diplomonads are small, mostly parasitic flagellates 542 Parabasilids are anaerobic endosymbionts that live in animals 542 Euglenoids and trypanosomes have both free-living species and parasites 543 Chromalveolates 544 Most dinoflagellates are a part of marine plankton 545 Apicomplexans are spore-forming parasites of animals 545 Ciliates use cilia for locomotion 547 Water molds produce biflagellate reproductive cells 548 Diatoms have shells composed of two parts 549 Brown algae are multicellular stramenopiles 549 Most golden algae are unicellular biflagellates 550 Rhizarians 551 Forams extend cytoplasmic projections that form a threadlike, interconnected net 551 Actinopods project slender axopods 552

Archaeplastids 552 Red algae do not produce motile cells 552 Green algae share many similarities with land plants 553 Unikonts 554 Amoebozoa are unikonts with lobose pseudopodia 555

27

Seedless Plants

The Evolution of Seed Plants 595 Our understanding of the evolution of flowering plants has made great progress in recent years 595

29

561

Fungal Reproduction 603 Many fungi reproduce asexually 603 Most fungi reproduce sexually 604

Bryophytes 565 Moss gametophytes are differentiated into “leaves” and “stems” 565 Liverwort gametophytes are either thalloid or leafy 567 Hornwort gametophytes are inconspicuous thalloid plants 570 Bryophytes are used for experimental studies 570 Recap: details of bryophyte evolution are based on fossils and on structural and molecular evidence 570

Fungal Diversity 605 Fungi are assigned to the opisthokont clade 605 Diverse groups of fungi have evolved 605 Chytrids have flagellate spores 606 Zygomycetes reproduce sexually by forming zygospores 607 Microsporidia have been a taxonomic mystery 609 Glomeromycetes are symbionts with plant roots 609 Ascomycetes reproduce sexually by forming ascospores 611 Basidiomycetes reproduce sexually by forming basidiospores 612

Seedless Vascular Plants 571 Club mosses are small plants with rhizomes and short, erect branches 572 Ferns are a diverse group of spore-forming vascular plants 572

Ecological Importance of Fungi 615 Fungi form symbiotic relationships with some animals 615 Mycorrhizae are symbiotic associations between fungi and plant roots 617 A lichen consists of two components: a fungus and a photoautotroph 617

Inquiring About Ancient Plants and Coal Formation 574 Some ferns and club mosses are heterosporous 576 Seedless vascular plants are used for experimental studies 578 Seedless vascular plants arose more than 420 mya 578

Seed Plants

Economic, Biological, and Medical Impact of Fungi 619 Fungi provide beverages and food 619 Fungi are important to modern biology and medicine 620 Fungi are used in bioremediation and to biologically control pests 621 Some fungi cause diseases in humans and other animals 621 Fungi cause many important plant diseases 622

582

An Introduction to Seed Plants

583

Gymnosperms 584 Conifers are woody plants that produce seeds in cones 584 Cycads have seed cones and compound leaves 587 Ginkgo biloba is the only living species in its phylum 587 Gnetophytes include three unusual genera 588 Flowering Plants 589 Monocots and eudicots are the two largest classes of flowering plants 590 Sexual reproduction takes place in flowers 590 The life cycle of flowering plants includes double fertilization 592 Seeds and fruits develop after fertilization 594 Flowering plants have many adaptations that account for their success 594 Floral structure provides insights into the evolutionary process 594

601

Characteristics of Fungi 602 Fungi absorb food from the environment 602 Fungi have cell walls that contain chitin 602 Most fungi consist of a network of filaments 602

Adaptations of Plants to Life on Land 562 The plant life cycle alternates between haploid and diploid generations 562 Four major groups of plants exist today 564

28

The Fungi

30

An Introduction to Animal Diversity Animal Characters

627

Adaptations to Ocean, Freshwater, and Terrestrial Habitats 627 Marine habitats offer many advantages 627 Some animals are adapted to freshwater habitats 628 Terrestrial living requires major adaptations 628 Animal Evolution 629 Molecular systematics helps biologists interpret the fossil record 629 Biologists develop hypotheses about the evolution of development 630

626

Reconstructing Animal Phylogeny 630 Animals exhibit two main types of body symmetry 630 Animal body plans are linked to the level of tissue development 631 Most bilateral animals have a body cavity lined with mesoderm 632 Bilateral animals form two main clades based on differences in development 633 Biologists have identified major animal clades based on structure, development, and molecular data 633

31

Jawless Fishes

Amniotes 693 Our understanding of amniote phylogeny is changing 693 Reptiles have many terrestrial adaptations 694 We can assign extant reptiles to five groups 695 Turtles have protective shells 696 Lizards and snakes are the most common modern reptiles 697 Tuataras superficially resemble lizards 697 Crocodilians have an elongated skull 697 How do we know that birds are really dinosaurs? 697 Modern birds are adapted for flight 699 Mammals have hair and mammary glands 700 New fossil discoveries are changing our understanding of the early evolution of mammals 700 Modern mammals are assigned to three subclasses 701

Sponges, Cnidarians, Ctenophores, and Protostomes 640 Sponges, Cnidarians, and Ctenophores: Animals with Asymmetry, Radial, or Biradial Symmetry 641 Sponges have collar cells and other specialized cells 641 Cnidarians have unique stinging cells 643 Comb jellies have adhesive glue cells that trap prey 647 The Lophotrochozoa 648 Flatworms are bilateral acoelomates 648 Nemerteans are characterized by their proboscis 651 Mollusks have a muscular foot, visceral mass, and mantle 652 Annelids are segmented worms 656 The lophophorates are distinguished by a ciliated ring of tentacles 658 Rotifers have a crown of cilia 660 The Ecdysozoa 661 Roundworms are of great ecological importance 661 Arthropods are characterized by jointed appendages and an exoskeleton of chitin 661

32

The Deuterostomes What Are Deuterostomes?

Part 6 STRUCTURE AND LIFE PROCESSES IN PLANTS 708 33

676

Echinoderms 676 Feather stars and sea lilies are suspension feeders 678 Many sea stars capture prey 678 Basket stars and brittle stars make up the largest group of echinoderms 679 Sea urchins and sand dollars have movable spines 679 Sea cucumbers are elongated, sluggish animals 679

Plant Meristems 718 Primary growth takes place at apical meristems 719 Secondary growth takes place at lateral meristems 719 Development of Form 720 The plane and symmetry of cell division affect plant form 721 The orientation of cellulose microfibrils affects the direction of cell expansion 721 Cell differentiation depends in part on a cell’s location 722 Morphogenesis occurs through pattern formation 724

680

Invertebrate Chordates 681 Tunicates are common marine animals 681 Lancelets clearly exhibit chordate characteristics 682 Systematists debate chordate phylogeny 682 Introducing the Vertebrates 683 The vertebral column is a derived vertebrate character 683 Vertebrate taxonomy is a work in progress 685

Plant Structure, Growth, and Development 708 The Plant Body 709 The plant body consists of cells and tissues 709 The ground tissue system is composed of three simple tissues 709 The vascular tissue system consists of two complex tissues 714 The dermal tissue system consists of two complex tissues 717

675

The Chordates: Defining Characteristics

685

Evolution of Jaws and Limbs: Jawed Fishes and Amphibians 687 Most cartilaginous fishes inhabit marine environments 687 The ray-finned fishes gave rise to modern bony fishes 689 Descendants of the lungfishes moved onto the land 690 Amphibians were the first successful land vertebrates 692

34

Leaf Structure and Function

728

Leaf Form and Structure 729 Leaf structure is adapted for maximum light absorption 730

Stomatal Opening and Closing 734 Blue light triggers stomatal opening 735 Additional factors affect stomatal opening and closing 736

37

Pollination 784 Many plants have mechanisms that prevent self-pollination 784 Flowering plants and their animal pollinators have coevolved 784 Some flowering plants depend on wind to disperse pollen 789

Leaf Abscission 738 In many leaves, abscission occurs at an abscission zone near the base of the petiole 738 Modified Leaves 738 Modified leaves of carnivorous plants capture insects 739

Stem Structure and Transport

Fertilization and Seed/Fruit Development 789 A unique double fertilization process occurs in flowering plants 789 Embryonic development in seeds is orderly and predictable 790 The mature seed contains an embryonic plant and storage materials 790 Fruits are mature, ripened ovaries 793 Seed dispersal is highly varied 794

744

Stem Growth and Structure 745 Herbaceous eudicot and monocot stems differ in internal structure 745 Woody plants have stems with secondary growth 746

36

Water Transport 753 Water and minerals are transported in xylem 753 Water movement can be explained by a difference in water potential 754 According to the tension–cohesion model, water is pulled up a stem 754 Root pressure pushes water from the root up a stem 756

Germination and Early Growth 796 Some seeds do not germinate immediately 796 Eudicots and monocots exhibit characteristic patterns of early growth 797

Translocation of Sugar in Solution 756 The pressure–flow model explains translocation in phloem 756

A Comparison of Sexual and Asexual Reproduction 799 Sexual reproduction has some disadvantages 800

Roots and Mineral Nutrition

781

The Flowering Plant Life Cycle 782 Flowers develop at apical meristems 782 Each part of a flower has a specific function 782

Transpiration and Guttation 736 Some plants exude liquid water 737

35

Reproduction in Flowering Plants

Asexual Reproduction in Flowering Plants Apomixis is the production of seeds without the sexual process 799

797

761

Root Structure and Function 762 Roots have root caps and root hairs 762 The arrangement of vascular tissues distinguishes the roots of herbaceous eudicots and monocots 763 Woody plants have roots with secondary growth 767 Some roots are specialized for unusual functions 767 Root Associations and Interactions 768 Mycorrhizae facilitate the uptake of essential minerals by roots 770 Rhizobial bacteria fix nitrogen in the roots of leguminous plants 771 The Soil Environment 772 Soil is composed of inorganic minerals, organic matter, air, and water 773 Soil organisms form a complex ecosystem 774 Soil pH affects soil characteristics and plant growth 775 Soil provides most of the minerals found in plants 775 Soil can be damaged by human mismanagement 776

38

Plant Developmental Responses to External and Internal Signals 803 Tropisms

804

Plant Hormones and Development 805 Plant hormones act by signal transduction 805 Auxins promote cell elongation 807 Gibberellins promote stem elongation 809 Cytokinins promote cell division 810 Ethylene promotes abscission and fruit ripening 810 Abscisic acid promotes seed dormancy 811 Brassinosteroids are plant steroid hormones 812 Identification of a universal flower-promoting signal remains elusive 812 Light Signals and Plant Development 813 Phytochrome detects day length 814 Competition for sunlight among shade-avoiding plants involves phytochrome 815 Phytochrome is involved in other responses to light, including germination 816 Phytochrome acts by signal transduction 816 Light influences circadian rhythms 816

ATP powers muscle contraction 852 The type of muscle fibers determines strength and endurance 855 Several factors influence the strength of muscle contraction 855 Smooth muscle and cardiac muscle are involuntary 856

Responses to Herbivores and Pathogens 817 Jasmonic acid activates several plant defenses 817 Methyl salicylate may induce systemic acquired resistance 818

Part 7 STRUCTURE AND LIFE PROCESSES IN ANIMALS 821

41

Neural Signaling

860

Neural Signaling: An Overview

39

Animal Structure and Function: An Introduction 821

Neurons and Glial Cells 862 Neurons receive stimuli and transmit neural signals 862 Certain regions of the CNS produce new neurons 862 Axons aggregate to form nerves and tracts 863 Glial cells play critical roles in neural function 863

Tissues, Organs, and Organ Systems 822 Epithelial tissues cover the body and line its cavities 822 Connective tissues support other body structures 823 Muscle tissue is specialized to contract 828 Nervous tissue controls muscles and glands 829 Tissues and organs make up the organ systems of the body 830 Inquiring About Unwelcome Tissues: Cancers

Transmitting Information along the Neuron 865 Ion channels and pumps maintain the resting potential of the neuron 865 Graded local signals vary in magnitude 867 Axons transmit signals called action potentials 867 The action potential is an all-or-none response 869 An action potential is self-propagating 870

831

Regulating the Internal Environment 834 Negative feedback systems restore homeostasis 834 A few positive feedback systems operate in the body 835

Transmitting Information across Synapses 872 Signals across synapses can be electrical or chemical 872 Neurons use neurotransmitters to signal other cells 872

Thermoregulation 835 Ectotherms absorb heat from their surroundings 836 Endotherms derive heat from metabolic processes 836 Many animals adjust to challenging temperature changes 837

40

Protection, Support, and Movement

Inquiring About Alzheimer’s Disease 874 Neurotransmitters bind with receptors on postsynaptic cells 875 Activated receptors can send excitatory or inhibitory signals 875

842

Neural Integration 877 Postsynaptic potentials are summed over time and space 877 Where does neural integration take place? 878

Epithelial Coverings 843 Invertebrate epithelium may secrete a cuticle 843 Vertebrate skin functions in protection and temperature regulation 843 Skeletal Systems 844 In hydrostatic skeletons, body fluids transmit force 844 Mollusks and arthropods have nonliving exoskeletons 845 Internal skeletons are capable of growth 845 The vertebrate skeleton has two main divisions 846 A typical long bone amplifies the motion generated by muscles 846 Bones are remodeled throughout life 847 Joints are junctions between bones 847 Muscle Contraction 848 Invertebrate muscle varies among groups 848 Insect flight muscles are adapted for rapid contraction 848 Vertebrate skeletal muscles act antagonistically to one another 849 A vertebrate muscle may consist of thousands of muscle fibers 850 Contraction occurs when actin and myosin filaments slide past one another 852

861

Neural Circuits: Complex Information Signaling

42

Neural Regulation

878

882

Invertebrate Nervous Systems: Trends in Evolution 883 The Vertebrate Nervous System: Structure and Function 884 Evolution of the Vertebrate Brain 886 The hindbrain develops into the medulla, pons, and cerebellum 886 The midbrain is prominent in fishes and amphibians 888 The forebrain gives rise to the thalamus, hypothalamus, and cerebrum 888 The Human Central Nervous System 888 The spinal cord transmits impulses to and from the brain 889 The most prominent part of the human brain is the cerebrum 891

The body follows a circadian cycle of sleep and wakefulness 894 The limbic system affects emotional aspects of behavior 897

44

937

Types of Circulatory Systems 938 Many invertebrates have an open circulatory system 939 Some invertebrates have a closed circulatory system 939 Vertebrates have a closed circulatory system 940

Inquiring About The Neurobiology of Traumatic Experience 898 Learning and memory involve long-term changes at synapses 899 Language involves comprehension and expression 901 The Peripheral Nervous System 902 The somatic division helps the body adjust to the external environment 902 The autonomic division regulates the internal environment 902

Vertebrate Blood 940 Plasma is the fluid component of blood 940 Red blood cells transport oxygen 941 White blood cells defend the body against disease organisms 942 Platelets function in blood clotting 943

Effects of Drugs on the Nervous System

Vertebrate Blood Vessels

904

Sensory Systems

The Human Heart 946 Each heartbeat is initiated by a pacemaker 948 The cardiac cycle consists of alternating periods of contraction and relaxation 949 The nervous system regulates heart rate 950 Stroke volume depends on venous return 951 Cardiac output varies with the body’s need 951

911

How Sensory Systems Work 912 Sensory receptors receive information 912 Sensory receptors transduce energy 912 Sensory input is integrated at many levels 912 We can classify sensory receptors based on location of stimuli or on the type of energy they transduce 914 Thermoreceptors

Nociceptors

Blood Pressure 951 Blood pressure varies in different blood vessels 953 Blood pressure is carefully regulated 954

915

Electroreceptors and Electromagnetic Receptors

The Pattern of Circulation 954 The pulmonary circulation oxygenates the blood 955 The systemic circulation delivers blood to the tissues 955

916

916

Mechanoreceptors 916 Tactile receptors are located in the skin 916 Proprioceptors help coordinate muscle movement 918 Many invertebrates have gravity receptors called statocysts 918 Hair cells are characterized by stereocilia 919 Lateral line organs supplement vision in fishes 919 The vestibular apparatus maintains equilibrium 919 Auditory receptors are located in the cochlea 920 Chemoreceptors 924 Taste receptors detect dissolved food molecules 925 The olfactory epithelium is responsible for the sense of smell 926 Many animals communicate with pheromones 926 Photoreceptors 927 Invertebrate photoreceptors include eyespots, simple eyes, and compound eyes 927 Vertebrate eyes form sharp images 928 The retina contains light-sensitive rods and cones 929 Light activates rhodopsin 930 Color vision depends on three types of cones 932 Integration of visual information begins in the retina 933

945

Evolution of the Vertebrate Cardiovascular System 946

Inquiring About Alcohol: The Most Abused Drug 905

43

Internal Transport

The Lymphatic System 956 The lymphatic system consists of lymphatic vessels and lymph tissue 956 The lymphatic system plays an important role in fluid homeostasis 957 Cardiovascular Disease

45

957

The Immune System: Internal Defense

963

Evolution of Immune Responses 964 Invertebrates launch nonspecific immune responses 964 Vertebrates launch nonspecific and specific immune responses 965 Nonspecific Immune Responses 966 Physical barriers prevent most pathogens from entering the body 966 Pattern recognition receptors activate nonspecific immune responses 966 Cells of the nonspecific immune system destroy pathogens and produce chemicals 966 Cytokines and complement mediate immune responses 967 Inflammation is a protective response 968

Specific Immune Responses 970 Many types of cells are involved in specific immune responses 971 The major histocompatibility complex is responsible for recognition of self 973 Cell-Mediated Immunity

47

973

The Vertebrate Digestive System 1016 Food processing begins in the mouth 1017 The pharynx and esophagus conduct food to the stomach 1017 Food is mechanically and enzymatically digested in the stomach 1018 Most enzymatic digestion takes place in the small intestine 1020 The liver secretes bile 1021 The pancreas secretes digestive enzymes 1021 Nutrients are digested as they move through the digestive tract 1021 Nerves and hormones regulate digestion 1022 Absorption takes place mainly through the villi of the small intestine 1023 The large intestine eliminates waste 1023

Response to Disease, Immune Failures, and Harmful Reactions 982 Cancer cells evade the immune system 982 Immunodeficiency disease can be inherited or acquired 983 HIV is the major cause of acquired immunodeficiency in adults 984 In an autoimmune disease, the body attacks its own tissues 986 Rh incompatibility can result in hypersensitivity 986 Allergic reactions are directed against ordinary environmental antigens 987 Graft rejection is an immune response against transplanted tissue 988

Gas Exchange

Required Nutrients 1024 Carbohydrates provide energy 1024 Lipids provide energy and are used to make biological molecules 1024 Proteins serve as enzymes and as structural components of cells 1026 Vitamins are organic compounds essential for normal metabolism 1026 Minerals are inorganic nutrients 1026 Antioxidants protect against oxidants 1027 Phytochemicals play important roles in maintaining health 1028

993

Adaptations for Gas Exchange in Air or Water

994

Types of Respiratory Surfaces 994 The body surface may be adapted for gas exchange 994 Tracheal tube systems deliver air directly to the cells 994 Gills are the respiratory surfaces in many aquatic animals 996 Terrestrial vertebrates exchange gases through lungs 996 The Mammalian Respiratory System 998 The airway conducts air into the lungs 998 Gas exchange occurs in the alveoli of the lungs 1001 Ventilation is accomplished by breathing 1001 The quantity of respired air can be measured 1002 Gas exchange takes place in the alveoli 1002 Gas exchange takes place in the tissues 1003 Respiratory pigments increase capacity for oxygen transport 1003 Carbon dioxide is transported mainly as bicarbonate ions 1004 Breathing is regulated by respiratory centers in the brain 1004 Hyperventilation reduces carbon dioxide concentration 1006 High flying or deep diving can disrupt homeostasis 1006 Some mammals are adapted for diving 1006 Breathing Polluted Air

1007

Inquiring About The Effects of Smoking

1012

Nutritional Styles and Adaptations 1013 Animals are adapted to their mode of nutrition 1013 Some invertebrates have a digestive cavity with a single opening 1015 Most animal digestive systems have two openings 1015

Antibody-Mediated Immunity 975 A typical antibody consists of four polypeptide chains 977 Antibodies are grouped in five classes 978 Antigen–antibody binding activates other defenses 978 The immune system responds to millions of different antigens 978 Monoclonal antibodies are highly specific 979 Immunological memory is responsible for long-term immunity 979

46

Processing Food and Nutrition

1008

Energy Metabolism 1029 Energy metabolism is regulated by complex signaling 1029 Obesity is a serious nutritional problem 1029 Undernutrition can cause serious health problems 1029

48

Osmoregulation and Disposal of Metabolic Wastes 1034 Maintaining Fluid and Electrolyte Balance Metabolic Waste Products

1035

1035

Osmoregulation and Excretion in Invertebrates 1036 Nephridial organs are specialized for osmoregulation and/or excretion 1037 Malpighian tubules conserve water 1038 Osmoregulation and Excretion in Vertebrates Freshwater vertebrates must rid themselves of excess water 1038 Marine vertebrates must replace lost fluid 1039 Terrestrial vertebrates must conserve water 1039

1038

The penis transfers sperm to the female 1083 Testosterone has multiple effects 1083 The hypothalamus, pituitary gland, and testes regulate male reproduction 1084

The Urinary System 1040 The nephron is the functional unit of the kidney 1042 Urine is produced by filtration, reabsorption, and secretion 1043 Urine becomes concentrated as it passes through the renal tubule 1045 Urine consists of water, nitrogenous wastes, and salts 1046 Hormones regulate kidney function 1046

49

Endocrine Regulation

Human Reproduction: The Female 1084 The ovaries produce gametes and sex hormones 1085 The oviducts transport the secondary oocyte 1086 The uterus incubates the embryo 1087 The vagina receives sperm 1087 The vulva are external genital structures 1088 The breasts function in lactation 1088 The hypothalamus, pituitary gland, and ovaries regulate female reproduction 1089

1052

An Overview of Endocrine Regulation 1053 The endocrine system and nervous system interact to regulate the body 1053 Negative feedback systems regulate endocrine activity 1053 Hormones are assigned to four chemical groups 1054

Inquiring About Breast Cancer 1090 Menstrual cycles stop at menopause 1093 Most mammals have estrous cycles 1093

Types of Endocrine Signaling 1055 Neurohormones are transported in the blood 1055 Some local regulators are considered hormones 1055

Fertilization, Pregnancy, and Birth 1093 Fertilization is the fusion of sperm and egg 1093 Inquiring About Novel Origins 1094 Hormones are necessary to maintain pregnancy 1095 The birth process depends on a positive feedback system 1096

Inquiring About Anabolic Steroids and Other Abused Hormones 1056 Mechanisms of Hormone Action 1057 Some hormones enter target cells and activate genes 1057 Many hormones bind to cell-surface receptors 1058 Neuroendocrine Regulation in Invertebrates

Human Sexual Response

Birth Control Methods and Abortion 1098 Many birth control methods are available 1098 Most hormone contraceptives prevent ovulation 1098 Intrauterine devices are widely used 1099 Barrier methods of contraception include the diaphragm and condom 1100 Emergency contraception is available 1100 Sterilization renders an individual incapable of producing offspring 1100 Future contraceptives may control regulatory peptides 1101 Abortions can be spontaneous or induced 1101

1060

Endocrine Regulation in Vertebrates 1060 Homeostasis depends on normal concentrations of hormones 1061 The hypothalamus regulates the pituitary gland 1061 The posterior lobe of the pituitary gland releases hormones produced by the hypothalamus 1061 The anterior lobe of the pituitary gland regulates growth and other endocrine glands 1062 Thyroid hormones increase metabolic rate 1066 The parathyroid glands regulate calcium concentration 1067 The islets of the pancreas regulate glucose concentration 1067 The adrenal glands help the body respond to stress 1071 Many other hormones are known 1073

Sexually Transmitted Diseases

51

Animal Development Development of Form

50

1097

1101

1106

1107

Asexual and Sexual Reproduction 1078 Asexual reproduction is an efficient strategy 1078 Most animals reproduce sexually 1078 Sexual reproduction increases genetic variability 1079

Fertilization 1107 The first step in fertilization involves contact and recognition 1107 Sperm entry is regulated 1108 Fertilization activates the egg 1109 Sperm and egg pronuclei fuse, restoring the diploid state 1109

Human Reproduction: The Male 1080 The testes produce gametes and hormones 1080 A series of ducts store and transport sperm 1081 The accessory glands produce the fluid portion of semen 1082

Cleavage 1109 The pattern of cleavage is affected by yolk 1109 Cleavage may distribute developmental determinants 1111 Cleavage provides building blocks for development 1112

Reproduction

1077

Gastrulation 1112 The amount of yolk affects the pattern of gastrulation 1113 Organogenesis

Culture in Vertebrate Societies 1148 Some vertebrates transmit culture 1148 Sociobiology explains human social behavior in terms of adaptation 1149

1115

Extraembryonic Membranes

1116

Human Development 1117 The placenta is an organ of exchange 1118 Organ development begins during the first trimester 1118 Development continues during the second and third trimesters 1120 More than one mechanism can lead to a multiple birth 1121 Environmental factors affect the embryo 1121 The neonate must adapt to its new environment 1121 Aging is not a uniform process 1123 Homeostatic response to stress decreases during aging 1123

52

Animal Behavior

Part 8 THE INTERACTIONS OF LIFE: ECOLOGY 1153 53

Features of Populations 1154 Density and dispersion are important features of populations 1154 Changes in Population Size 1156 Dispersal affects the growth rate in some populations 1156 Each population has a characteristic intrinsic rate of increase 1157 No population can increase exponentially indefinitely 1157

1127

Behavior and Adaptation 1128 Behaviors have benefits and costs 1128 Genes interact with environment 1128 Behavior depends on physiological readiness 1130 Many behavior patterns depend on motor programs 1130

Factors Influencing Population Size 1158 Density-dependent factors regulate population size 1158 Density-independent factors are generally abiotic 1160

Learning: Changing Behavior as a Result of Experience 1131 An animal habituates to irrelevant stimuli 1131 Imprinting occurs during an early critical period 1131 In classical conditioning, a reflex becomes associated with a new stimulus 1132 In operant conditioning, spontaneous behavior is reinforced 1132 Animal cognition is controversial 1133 Play may be practice behavior 1134 Behavioral Responses to Environmental Stimuli Biological rhythms regulate many behaviors 1134 Environmental signals trigger physiological responses that lead to migration 1135 Foraging Behavior

Introduction to Ecology: Population Ecology 1153

Life History Traits 1161 Life tables and survivorship curves indicate mortality and survival 1163 Metapopulations

Human Populations 1165 Not all countries have the same growth rate 1167 The age structure of a country helps predict future population growth 1168 Environmental degradation is related to population growth and resource consumption 1169

1134

1136

Costs and Benefits of Social Behavior 1137 Communication is necessary for social behavior 1138 Dominance hierarchies establish social status 1139 Many animals defend a territory 1140 Some insect societies are highly organized 1141 Sexual Selection 1142 Animals of the same sex compete for mates 1142 Animals select quality mates 1143 Sexual selection favors polygynous mating systems 1143 Some animals care for their young 1144 Helping Behavior 1146 Altruistic behavior can be explained by inclusive fitness 1146 Helping behavior may have alternative explanations 1147 Some animals help nonrelatives 1148

1164

54

Community Ecology

1173

Community Structure and Functioning 1174 Community interactions are complex and often not readily apparent 1175 The niche is a species’ ecological role in the community 1175 Competition is intraspecific or interspecific 1177 Natural selection shapes the bodies and behaviors of both predator and prey 1180 Symbiosis involves a close association between species 1182 Inquiring About Mimicry in Butterflies

1183

Strength and Direction of Community interactions 1185 Other species of a community depend on or are greatly affected by keystone species 1185 Dominant species influence a community as a result of their greater size or abundance 1186 Ecosystem regulation occurs from the bottom up and top down 1186

Aquatic Ecosystems 1228 Freshwater ecosystems are linked to land and marine ecosystems 1228 Estuaries occur where fresh water and salt water meet 1231 Marine ecosystems dominate Earth’s surface 1232

Community Biodiversity 1187 Ecologists seek to explain why some communities have more species than others 1188 Species richness may promote community stability 1190 Community Development 1190 Disturbance influences succession and species richness 1191 Ecologists continue to study community structure 1192

55

Ecosystems and the Biosphere

Cycles of Matter in Ecosystems 1203 Carbon dioxide is the pivotal molecule in the carbon cycle 1203 Bacteria are essential to the nitrogen cycle 1204 The phosphorus cycle lacks a gaseous component 1206 Water moves among the ocean, land, and atmosphere in the hydrologic cycle 1207 1208

Inquiring About Life without the Sun 1209 The atmosphere contains several gases essential to organisms 1210 The global ocean covers most of Earth’s surface 1211 Climate profoundly affects organisms 1212 Fires are a common disturbance in some ecosystems 1213 Studying Ecosystem Processes

56

1236

Biogeography 1237 Land areas are divided into six biogeographic realms 1238

1196

Energy Flow through Ecosystems 1197 Ecological pyramids illustrate how ecosystems work 1199 Ecosystems vary in productivity 1200 Food chains and poisons in the environment 1202

Abiotic Factors in Ecosystems The sun warms Earth 1208

Ecotones

1214

Ecology and the Geography of Life

1218

Biomes 1219 Tundra is the cold, boggy plains of the far north 1219 Inquiring About The Distribution of Vegetation on Mountains 1220 Boreal forest is the evergreen forest of the north 1221 Temperate rain forest has cool weather, dense fog, and high precipitation 1221 Temperate deciduous forest has a canopy of broad-leaf trees 1222 Temperate grasslands occur in areas of moderate precipitation 1222 Chaparral is a thicket of evergreen shrubs and small trees 1223 Deserts are arid ecosystems 1224 Savanna is a tropical grassland with scattered trees 1225 There are two basic types of tropical forests 1225

57

Biological Diversity and Conservation Biology 1242 The Biodiversity Crisis 1243 Endangered species have certain characteristics in common 1244 Inquiring About Declining Amphibian Populations 1245 Human activities contribute to declining biological diversity 1246 Conservation Biology 1249 In situ conservation is the best way to preserve biological diversity 1249 Ex situ conservation attempts to save species on the brink of extinction 1251 The Endangered Species Act provides some legal protection for species and habitats 1252 International agreements provide some protection of species and habitats 1253 Deforestation 1253 Why are tropical rain forests disappearing? 1254 Why are boreal forests disappearing? 1255 Climate Change 1255 Greenhouse gases cause climate change 1256 What are the probable effects of climate change? 1257 Declining Stratospheric Ozone 1259 Certain chemicals destroy stratospheric ozone 1259 Ozone depletion harms organisms 1260 International cooperation is helping repair the ozone layer 1260 Appendix A Periodic Table of the Elements Appendix B Classification of Organisms

A-2

Appendix C Understanding Biological Terms Appendix D Abbreviations

A-1

A-6

A-9

Appendix E Answers to Test Your Understanding Questions A-11 Glossary Index

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Preface

This ninth edition of Solomon, Berg, Martin Biology conveys our vision of the dynamic science of biology and how it affects every aspect of our lives, from our health and behavior to the challenging environmental issues that confront us. New discoveries in the biological sciences continue to increase our understanding of both the unity and diversity of life’s processes and adaptations. With this understanding, we become ever more aware of our interdependence with the vast diversity of organisms with which we share planet Earth.

BIOLOGY: THE STUDENT-FRIENDLY BIOLOGY BOOK We want beginning students to experience learning biology as an exciting journey of discovery. In the ninth edition of Biology, we explore Earth’s diverse organisms, their remarkable adaptations to the environment, and their evolutionary and ecological relationships. We present the workings of science and the contributions of scientists whose discoveries not only expand our knowledge of biology but also help shape and protect the future of our planet. Biology provides insight into what science is, how scientists work, what scientists have contributed, and how scientific knowledge affects daily life. Since the first edition of Biology, we have worked to present the principles of biology in an integrated way that is accurate, interesting, and conceptually accessible to students. In this ninth edition of Biology, we continue this tradition. We also continue to present biology in an inquiry-based framework. Some professors interpret inquiry as a learning method that takes place in the laboratory as students perform experiments. Laboratory research is certainly an integral part of inquiry-based learning. But inquiry is also a way of learning in which the student actively pursues knowledge outside the laboratory. In Biology, we have always presented the history of scientific advances, including scientific debates, to help students understand that science is a process, that is, a field of investigative inquiry, as well as a body of knowledge (the product of inquiry). In the ninth edition of Biology, we make a concerted effort to further integrate inquiry-based learning into the textbook with the introduction of new features and the expansion of several others (discussed in the following sections). Throughout the text, we stimulate interest by relating concepts to experiences within the student’s frame of reference. By helping students make such connections, we facilitate their mastery of general concepts. We hope the combined effect of an engaging writing style and interesting features will motivate and excite students in their study of biology.

THE SOLOMON/BERG/MARTIN LEARNING SYSTEM In the ninth edition, we have continued to refine our highly successful Learning System. This system provides the student with the learning strategies needed to integrate biological concepts and to demonstrate mastery of these concepts. Learning biology is challenging because the subject of biology is filled with so many new terms and so many facts that must be integrated into the framework of general biological principles. To help students focus on important principles and concepts, we provide Learning Outcomes for the course and Learning Objectives for each major section of every chapter. At the end of each section, we provide Review Questions based on the Learning Objectives so students can assess their level of understanding of the material presented in the section. At the end of each chapter, we include a Summary: Focus on Learning Objectives that is organized around the Learning Objectives. The Summary is followed by Test Your Understanding questions (including multiple-choice exercises and diagram labeling) and Critical Thinking questions. Throughout the book, students are directed to CengageNOW, a powerful online diagnostic tool that helps students assess their study needs and master the chapter objectives. After taking a pretest on CengageNOW, students receive feedback based on their answers as well as a Personalized Study Plan with links to animations and other resources keyed to their specific learning needs. Selected illustrations in the text are also keyed to Animated figures in CengageNOW.

Pedagogical Features Our Learning System includes numerous learning strategies that help students increase their success: •

NEW An updated and expanded art program reinforces concepts discussed in the text, and presents complex processes in clear steps. This edition expands the number of Key Experiment figures, which encourage students to evaluate investigative approaches that scientists have taken. Key Experiment figures emphasize the scientific process in both classic and modern research; examples include Figures 8-9, 20-13, 22-9, 27-11, and 52-21. Also expanded in this edition are Key Point figures, in which important concepts are stated in process diagrams of complex topics; examples include Figures 1-11, 6-6, 8-2, 29-4, and 32-7. Many of the Key Point figures have numbered parts that show sequences of events in biological processes or life cycles. Numerous photographs, both alone

and combined with line art, help students grasp concepts by connecting the “real” to the “ideal.” The line art uses features such as orientation icons to help the student put the detailed figures into the broader context. We use symbols and colors consistently throughout the book to help students connect concepts. For example, the same four colors and shapes are used throughout the book to identify guanine, cytosine, adenine, and thymine. Similarly, the same colors are used consistently in illustrations and tables to indicate specific clades of organisms. •

NEW Research Method figures describe why biologists use a particular method and explain how the method is executed. Examples include Figures 12-6, 15-5, and 23-11.

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knowledge. Answers to the Test Your Understanding questions are provided in Appendix E. Thought-provoking Critical Thinking questions encourage the student to apply the concepts just learned to new situations or to make connections among important concepts. Each chapter has one or more Evolution Link questions in the Critical Thinking section. Many chapters contain one or more Analyzing Data questions that require students to actively interpret experimental data presented in the chapter. NEW Many Critical Thinking sections also include one or more Science, Technology, and Society questions. The Glossary at the end of the book, the most comprehensive glossary found in any biology text, provides precise definitions of terms. The Glossary is especially useful because it is extensively cross-referenced and includes pronunciations for many terms. The vertical blue bar along the margin facilitates rapid access to the Glossary. The companion website also includes glossary flash cards with audio pronunciations.

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NEW Inquiring About boxes explore issues of special relevance to students, such as the effects of smoking, how traumatic experiences affect the body, and breast cancer. These boxes also provide a forum for discussing some interesting topics in more detail, such as the smallest ancient humans, ancient plants and coal formation, hydrothermal vent communities, and declining amphibian populations.

Course Learning Outcomes

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A list of Key Concepts at the beginning of each chapter provides a chapter overview and helps the student focus on important principles discussed in the chapter.

At the end of a successful study of introductory biology, the student can demonstrate mastery of biological concepts by responding accurately to the following Course Learning Outcomes:

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Learning Objectives at the beginning of each major section in the chapter indicate, in behavioral terms, what the student must do to demonstrate mastery of the material in that section.

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Each major section of the chapter is followed by a series of Review questions that assess comprehension by asking the student to describe, explain, compare, contrast, or illustrate important concepts. The Review questions are based on the section Learning Objectives.

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Concept Statement Subheads introduce sections, previewing and summarizing the key idea or ideas to be discussed in that section. Sequence Summaries within the text simplify and summarize information presented in paragraph form. For example, paragraphs describing blood circulation through the body or the steps by which cells take in certain materials are followed by a Sequence Summary listing the sequence of structures or steps. Numerous tables, many illustrated, help the student organize and summarize material presented in the text. Many tables are color-coded. A Summary: Focus on Learning Objectives at the end of each chapter is organized around the chapter Learning Objectives. This summary provides a review of the material, and because selected key terms are boldfaced in the summary, students learn vocabulary words within the context of related concepts. End-of-chapter questions provide students with the opportunity to evaluate their understanding of the material in the chapter. Test Your Understanding consists of multiple-choice questions, some of which are based on the recall of important terms, whereas others challenge students to integrate their

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Design an experiment to test a given hypothesis, using the procedure and terminology of the scientific method. Cite the cell theory, and relate the structure of organelles to their functions in both prokaryotic and eukaryotic cells. Describe the mechanisms of evolution, explain why evolution is the principal unifying concept in biology, and discuss natural selection as the primary agent of evolutionary change. Explain the role of genetic information in all species, and discuss applications of genetics that affect society. Describe several mechanisms by which cells and organisms transfer information, including the use of nucleic acids in genetic transmission of information, signal transduction, chemical signals (such as hormones and pheromones), electrical signals (for example, neural transmission), sounds, and visual displays. Argue for or against the classification of organisms in three domains and several kingdoms or supergroups, characterizing each of these clades; based on your knowledge of genetics and evolution, give specific examples of the unity and diversity of organisms in different domains and supergroups. Compare the structural adaptations, life processes, and life cycles of a prokaryote, protist, fungus, plant, and animal. Define homeostasis, and give examples of regulatory mechanisms, including feedback systems. Trace the flow of matter and energy through a photosynthetic cell and a nonphotosynthetic cell, and through the biosphere, comparing the roles of producers, consumers, and decomposers. Describe the study of ecology at the levels of an individual organism, a population, a community, and an ecosystem.

WHAT’S NEW: AN OVERVIEW OF BIOLOGY, NINTH EDITION Three themes are interwoven throughout Biology: the evolution of life, the transmission of biological information, and the flow of energy through living systems. As we introduce the concepts of modern biology, we explain how these themes are connected and how life depends on them. Educators present the major topics of an introductory biology course in a variety of orders. For this reason, we carefully designed the eight parts of this book so that they do not depend heavily on preceding chapters and parts. This flexible organization means that an instructor can present the 57 chapters in any number of sequences with pedagogical success. Chapter 1, which introduces the student to the major principles of biology, provides a comprehensive springboard for future discussions, whether the professor prefers a “top-down” or “bottom-up” approach. In this edition, as in previous editions, we examined every line of every chapter for accuracy and currency, and we made a careful attempt to update every topic and verify all new material. The following brief summary provides a general overview of the organization of Biology and some changes made to the ninth edition.

Part 1 The Organization of Life The six chapters that make up Part 1 provide basic principles of biology and the concepts of chemistry and cell biology that lay the foundation upon which the remaining parts of the book build. We begin Chapter 1 with a discussion of influenza pandemics and how scientists respond to new varieties of pathogens. We then introduce the main themes of the book—evolution, information transfer, and energy transfer. Chapter 1 examines several fundamental concepts in biology and the nature of the scientific process, including a discussion of systems biology. Chapters 2 and 3, which focus on the molecular level of organization, establish the foundations in chemistry necessary for understanding biological processes. Chapters 4, 5, and 6 focus on the cellular level of organization, including cell structure and function, cell membranes, and cell signaling. We have expanded coverage of signaling pathways and the evolution of cell communication. For example, we added a discussion of Ras pathways, as well as a Key Experiment on a Ras pathway that regulates reproduction in mammals.

Part 2 Energy Transfer through Living Systems Because all living cells need energy for life processes, the flow of energy through living systems—that is, capturing energy and converting it to usable forms—is a basic theme of Biology. Chapter 7 examines how cells capture, transfer, store, and use energy. Chapters 8 and 9 discuss the metabolic adaptations by which organisms obtain and use energy through cellular respiration and photosynthesis. This section has a new Key Experiment figure on the evidence for chemiosmosis. Also, the concept of resonance has been included in the explanation for energy transfer within a photosystem.

Part 3 The Continuity of Life: Genetics We have updated and expanded the eight chapters of Part 3 for the ninth edition. We begin this unit by discussing mitosis and meiosis in Chapter 10, which includes a new Key Experiment on using laser photobleaching to determine how chromosomes are transported toward the spindle poles during anaphase and a new figure on key checkpoints in the cell cycle. Chapter 11, which considers Mendelian genetics and related patterns of inheritance, has new paragraphs on nature versus nurture in human characters and on hydrangea flower color as an illustration of the influence of the environment on the phenotype. We then turn our attention to the structure and replication of DNA in Chapter 12. We include new photographs of Rosalind Franklin, Watson and Crick, phages, DNA replication in E. coli, and telomeres. Chapter 13 has a discussion of RNA and protein synthesis, including new information on the contributions of Roger Kornberg, transposons in the human genome, and the percentage of DNA coding for polypeptides versus the percentage of our genome that is expressed. Gene regulation is discussed in Chapter 14, which includes new material on eukaryotic promoters and enhancers and silencers, and on epigenetic inheritance. In Chapter 15, we focus on DNA technology and genomics, including an expanded discussion of engineered strains of E. coli, nucleic acid probes, Northern and Western blots, and molecular evolution. These chapters build the necessary foundation for exploring human genetics and the human genome in Chapter 16, which includes new sections on genomic imprinting and on genome-wide association (GWA) scans. In Chapter 17, we introduce the role of genes in development, emphasizing studies on specific model organisms that have led to spectacular advances in this field; changes include new material on induced pluripotent stem (iPS) cells.

Part 4 The Continuity of Life: Evolution Although we explore evolution as the cornerstone of biology throughout the book, Part 4 discusses evolutionary concepts in depth. We provide the history behind the discovery of the scientific theory of evolution, the mechanisms by which it occurs, and the methods by which it is studied and tested. Chapter 18 introduces the Darwinian concept of evolution and presents several kinds of evidence that support the scientific theory of evolution. In Chapter 19, we examine evolution at the population level. Chapter 20 describes the evolution of new species and discusses aspects of macroevolution. Chapter 21 summarizes the evolutionary history of life on Earth. In Chapter 22, we recount the evolution of the primates, including humans. Many topics and examples have been added to the ninth edition, such as new material on genome sequencing, whale evolution, genetic polymorphism, cichlid evolution, hybrid zones, the metabolism-first hypothesis, and recent discoveries in human evolution, including new material on Ardipithecus, Homo ergaster, archaic humans (H. antecessor and H. heidelbergensis), and Neandertals.

Part 5 The Diversity of Life Emphasizing the cladistic approach, we use an evolutionary framework to discuss each group of organisms. We present current

hypotheses of how groups of organisms are related. Chapter 23 discusses why organisms are classified and provides insight into the scientific process of deciding how they are classified. New advances have enabled us to further clarify the connection between evolutionary history and systematics in the ninth edition. NEW Chapter 24 now focuses entirely on the viruses. We added a discussion of polydnaviruses, illustrated by a new Key Experiment. We also expanded the discussion of subviral agents. NEW Chapter 25 is now devoted to the prokaryotes. We added information on the archaea, including a brief discussion of their flagella. The discussion of antibiotic resistance was expanded, and an image of methicillin-resistant Staphylococcus aureus (MRSA) was added. Chapter 26, which describes the protists, was completely reorganized and updated to reflect recent evolutionary data; a new colorcoded table summarizes the five “supergroups” of eukaryotes. Chapters 27 and 28 present the members of the plant kingdom; Chapter 28 includes new cladograms showing the evolution of land plants and the evolution of seedless vascular plants. Discussion of the origin and early evolution of angiosperms is included in Chapter 28. Chapter 29 describes the fungi. We have added a description of symbiotic relationships among fungi, grasses, and viruses and also a discussion of cell signaling in mycorrhizal fungi between fungi and root cells. In Chapters 30 through 32, we discuss the diversity of animals. We have reorganized some of the discussion and added several tables. We emphasize the cladistic approach, although we continue to include traditional phyla and classes.

Part 6 Structure and Life Processes in Plants Part 6 introduces students to the fascinating plant world. It stresses relationships between structure and function in plant cells, tissues, organs, and individual organisms. In Chapter 33, we introduce plant structure, growth, and differentiation. Chapter 33 contains a new section on plant form that includes plant aspects of cell division, cell expansion, cell differentiation, tissue culture, morphogenesis, pattern formation, positional information, and Arabidopsis mutants. Chapters 34 through 36 discuss the structural and physiological adaptations of leaves, stems, and roots; these chapters include new discussion of the molecular control of plant leaf form, cotransport in phloem loading, and transcription factors that affect root hair development. Chapter 37 describes reproduction in flowering plants, including asexual reproduction, flowers, fruits, and seeds. New to this edition is a spread of photos and a table showing floral characteristics associated with various animal pollinators. Chapter 38 focuses on growth responses and regulation of growth. In the ninth edition, we present the latest findings generated by the continuing explosion of knowledge in plant biology, particularly at the molecular level. New topics include advances in auxin signaling, GA signaling, cytokinin signaling, ethylene signaling, and ABA receptors.

Part 7 Structure and Life Processes in Animals In Part 7, we provide a strong emphasis on comparative animal physiology, showing the structural, functional, and behavioral adaptations that help animals meet environmental challenges. We use a comparative approach to examine how various animal groups have solved both similar and diverse problems. In Chapter 39, we

discuss the basic tissues and organ systems of the animal body, homeostasis, and the ways that animals regulate their body temperature. Chapter 40 focuses on different types of body coverings, skeletons, and muscles, and discusses how they function. In Chapters 41 through 43, we discuss neural signaling, neural regulation, and sensory reception. In Chapters 44 through 51, we compare how different animal groups carry on life processes, such as internal transport, internal defense, gas exchange, digestion, reproduction, and development. Each chapter in this part considers the human adaptations for the life processes being discussed. Part 7 ends with a discussion of behavioral adaptations in Chapter 52. Reflecting recent research findings, we have revised or added new material on many topics, including neurotransmitters, learning, memory, sleepwake cycles, coevolution of bats and moths, cardiovascular disease, Toll-like receptors, chronic inflammation, and certain hormones. The art program has been updated and improved, and many new photographs and photomicrographs have been added.

Part 8 The Interactions of Life: Ecology Part 8 focuses on the dynamics of populations, communities, and ecosystems and on the application of ecological principles to disciplines such as conservation biology. Chapters 53 through 56 give the student an understanding of the ecology of populations, communities, ecosystems, and the biosphere, whereas Chapter 57 focuses on global environmental issues. Among the many new topics in this unit are correlations between lemming populations and climate change; a 2006 study by Peter and Rosemary Grant on character displacement in ground finches; evolution in scarlet king snakes away from their model, the coral snake; Paine’s classic work with predatory Pisaster sea stars in rocky intertidal communities; ocean acidification; invasion of many of the world’s grasslands by woody trees and shrubs; and data on climate change from the Fourth IPCC Assessment.

A COMPREHENSIVE PACKAGE FOR LEARNING AND TEACHING A carefully designed supplement package is available to further facilitate learning. In addition to the usual print resources, we are pleased to present student multimedia tools that have been developed in conjunction with the text.

Resources for Students Aplia is an online interactive homework solution that stimulates active learning by increasing student effort and engagement. Aplia enables students to optimize the time they spend devoted to coursework. Aplia courses are customized to fit with each instructor’s syllabus, and provide automatically graded homework with detailed, immediate feedback on every question. Aplia’s interactive tools also serve to increase student engagement and understanding. The Aplia assignments allow students to apply what they learn in the text and in face-to-face instruction directly to their homework.

Aplia homework promotes active learning by: • • •

Leading students through the thought processes of doing science Helping students understand the scientific procedures that lead to scientific results covered in the textbook Connecting conceptual figures with scientific issues and problems

Aplia Gradebook Analytics show how each student is doing relative to peers in the class. This allows instructors to quickly and easily: • • •

Review progress student by student, and even drill down to the homework and question level Use graphs to monitor student, and class, performance Determine in what subject areas students are struggling and need more review

Straightforward language, a simple and intuitive user interface, and online material that reinforces the textbook all combine to make Aplia a solution that makes students think and helps students learn and retain knowledge. CengageNOW. This updated and expanded online homework and learning tool helps students assess personal study needs and focus their time. By taking a pretest, they are provided with a Personalized Study Plan that directs them to text sections and narrated animations—many new to this edition—that they need to review. If they need to brush up on basic skills, the How Do I Prepare? feature walks them through tutorials on basic math, chemistry, study skills, and word roots. Homework and tests can be assigned through CengageNOW, with automatic grading for all multiplechoice and true/false questions. Biology CourseMate. Cengage Learning’s Biology CourseMate brings course concepts to life with interactive learning, study, and exam preparation tools that support the printed textbook. Watch student comprehension soar as the class works with the printed textbook and the textbook-specific website. Biology CourseMate includes: • •

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An interactive eBook Interactive teaching and learning tools including: • Quizzes • Videos • Animations • Flash cards • And more Engagement Tracker, a first-of-its-kind tool that monitors student engagement in the course

Go to www.cengagebrain.com to access these resources, and look , which denotes a resource available within for this icon CourseMate. Study Guide to accompany Biology, Ninth Edition, by Jennifer Aline Metzler of Ball State University and Robert Yost of Indiana University and Purdue University, Indianapolis. Updated for this edition, the study guide provides the student with many opportunities to review chapter concepts. Multiple-choice study questions, coloring-book exercises, vocabulary-building exercises, and

many other types of active-learning tools are provided to suit different cognitive learning styles. A Problem-based Guide to Basic Genetics by Donald Cronkite of Hope College. This brief guide provides students with a systematic approach to solving genetics problems, along with numerous solved problems and practice problems. Spanish Glossary. This Spanish glossary of biology terms is available to Spanish-speaking students. Audio Study Tools. This edition of Biology is accompanied by useful study tools, which contain valuable information such as reviews of important concepts, key terms, questions, and study tips. Students can download the audio study tools. Virtual Biology Laboratory 4.0. Now with an upgraded user interface, these 14 online laboratory experiments allow students to “do” science by acquiring data, performing simulated experiments, and using data to explain biological concepts. Assigned activities automatically flow to the instructor’s grade book. Self-designed activities ask students to plan their procedures around an experimental question and write up their results.

Additional Resources for Instructors The instructors’ examination copy for this edition lists a comprehensive package of print and multimedia supplements, including online resources, available to qualified adopters. Please ask your local sales representative for details.

ACKNOWLEDGMENTS The development and production of the ninth edition of Biology required extensive interaction and cooperation among the authors and many individuals in our family, social, and professional environments. We thank our editors, colleagues, students, family, and friends for their help and support. Preparing a book of this complexity is challenging and requires a cohesive, talented, and hardworking professional team. We appreciate the contributions of everyone on the editorial and production staff at Brooks/Cole– Cengage Learning who worked on this ninth edition of Biology. We thank our Publisher Yolanda Cossio for her commitment to Biology and for working closely with us throughout the entire process of development and production. We appreciate the help of our Developmental Editor, Christopher Delgado, who diligently kept us on track and coordinated many aspects of this complex project. We thank Tom Ziolkowski, our Marketing Manager, whose expertise ensured that you would know about our new edition. We appreciate the help of Senior Content Project Manager Hal Humphrey, who guided overall production of the project. We thank Editorial Assistants Brandusa Radoias and Joshua Taylor for quickly providing us with resources whenever we needed them. We thank Creative Director Rob Hugel, Art Director and Cover Designer John Walker, and Text Designer tani hasegawa. We also thank Jean Thompson for developing the instructor’s preface. We appreciate the work of Lauren Oliveira, Media Editor, who coordinated the many high-tech components of the computerized

aspects of our Learning System. We thank Alexis Glubka, Assistant Editor, for coordinating the print supplements. We are grateful to our Production Editor, Joan Keyes of Dovetail Publishing Services, for coordinating the many editors involved in the preparation of this edition and bringing together the thousands of complex pieces of the project to produce Biology, Ninth Edition. We value the careful work of our Copy Editor, Susan Gall, who patiently helped us maintain consistency and improve the manuscript. We thank Amanda Bickel of Precision Graphics for helping us greatly improve the art program for this book. We also appreciate the efforts of Photo Editor Tim Herzog of Bill Smith Group in helping us find excellent images. We appreciate the help, patience, and hard work of our production team. Our schedule for this project was very demanding. At times, it seemed like the team worked around the clock. When we sent an email at 11 p.m. or during the weekend, we often received an immediate response. These dedicated professionals and many others on the Brooks/ Cole team provided the skill, attention, patience, and good humor needed to produce Biology, Ninth Edition. We thank them for their help and support throughout this project. We thank Dr. Susan Pross, University of South Florida, College of Medicine, for her helpful suggestions for updating the immunology chapter. We appreciate the help of obstetrician/gynecologist Dr. Amy Solomon for her input regarding pregnancy, childbirth, contraception, and sexually transmitted diseases. We appreciate the expert assistance of Mary Kay Hartung, Research and Information Specialist at Florida Gulf Coast University, who we counted on for help whenever we had difficulty finding needed research studies and other information. We thank doctoral student Lois Ball of the University of South Florida, who calculated scale bars for many of the photomicrographs. We are grateful to Mical Solomon for his computer help. We thank our families and friends for their understanding, support, and encouragement as we struggled through many revisions and intense deadlines. We especially thank Dr. Kathleen M. Heide, Freda Brod, Alan Berg, Jennifer and Pat Roath, Dr. Charles Martin, and Margaret Martin for their support and input. Our colleagues and students who have used our book have provided valuable input by sharing their responses to past editions of Biology. We thank them and ask again for their comments and suggestions as they use this new edition. We can be reached through the Internet via our website at www.CengageBrain.com or through our editors at Brooks/Cole, a division of Cengage Learning. We express our thanks to the many biologists who have read the manuscript during various stages of its development and provided us with valuable suggestions for improving it. Ninth edition reviewers include the following: Karl Aufderheide, Texas A & M University Felicitas Avendano, Grand View College, Iowa Brian Bagatto, University of Akron Catherine Black, Idaho State University Susan Boothe, Alabama Southern Community College Mark Browning, Purdue University Jill Buettner, Richland College Doug Burks, Wilmington College, Ohio

David Byres, Florida Community College at Jacksonville Geoffrey Church, Fairfield University Reggie Cobb, Nash Community College Barbara Collins, California Lutheran University Karen Curto, University of Pittsburgh Phillippa Drennan, Loyola Marymount University James DuMond, Texas Southern University Wayne Elisens, University of Oklahoma Cheryld Emmons, Alfred University, New York Suzanne Gollery, Sierra Nevada College Jack Holt, Susquehanna University Ursula Howson, Monmouth University Ken Klemow, Wilkes University Paula Lemons, Duke University Diana Lipscomb, George Washington University Christopher Loretz, University of Buffalo Kathi Maleug, University of Colorado at Colorado Springs Paul Manos, Duke University Helen McDearman, University of Tennessee at Chattanooga Virginia McDonough, Hope College Betsy Morgan, Lonestar College Dana Nayduch, Georgia Southern University Onesimus Otieno, Oakwood University Michael Pallidino, Monmouth College Craig Peebles, University of Pittsburgh Ed Perry, Faulkner State Community College Eric Ribbens, Western Illinois University Laurel Roberts, University of Pittsburgh Darrin Rubino, Hanover College, Indiana Timothy Schuh, St. Cloud State University Sharon Thoma, University of Wisconsin Joanne Tillotson, Purchase College, State University of New York Timothy Tripp, Sam Houston State University Paul Trombley, Florida State University Mary Tyler, University of Maine Fred Wasserman, Boston University Susan Whittemore, Keene State College Lawrence Williams, University of Houston Steven Wilt, Bellarmine College, Kentucky Robert Yost, Indiana University–Purdue University, Indianapolis Wendy Zomlefer, University of Georgia We would also like to thank the hundreds of previous edition reviewers, both professors and students, who are too numerous to mention. They asked thoughtful questions, provided new perspectives, offered alternative wordings to clarify difficult passages, and informed us of possible errors. We are truly indebted to their excellent feedback. Their suggestions have helped us improve each edition of Biology.

To the Student

Biology is a challenging subject. The thousands of students we have taught have differed in their life goals and learning styles. Some have had excellent backgrounds in science; others, poor ones. Regardless of their backgrounds, it is common for students taking their first college biology course to find that they must work harder than they expected. The Learning System we use in this book is described in the Preface. Using the strategies of the Learning System will help you master the language and concepts of biology. You will also want to use the many online tools available to Biology students. These tools, also described in the Preface, include Aplia, CengageNOW, and Biology CourseMate. In addition to these learning strategies, you can make the task of learning biology easier by using approaches that have been successful to a broad range of our students over the years.

Make a Study Schedule Many college professors suggest that students study 3 hours for every hour spent in class. This major investment in study time is one of the main differences between high school and college. To succeed academically, college students must learn to manage their time effectively. The actual number of hours you spend studying biology will vary depending on how quickly you learn the material, as well as on your course load and personal responsibilities, such as work schedules and family commitments. The most successful students are often those who are best organized. At the beginning of the semester, make a detailed daily calendar. Mark off the hours you are in each class, along with travel time to and from class if you are a commuter. After you get your course syllabi, add to your calendar the dates of all exams, quizzes, papers, and reports. As a reminder, it also helps to add an entry for each major exam or assignment 1 week before the test or due date. Now add your work schedule and other personal commitments to your calendar. Using a calendar helps you find convenient study times. Many of our successful biology students set aside 2 hours a day to study biology rather than depend on a weekly marathon session for 8 or 10 hours during the weekend (when that kind of session rarely happens). Put your study hours into your daily calendar, and stick to your schedule.

Determine Whether the Professor Emphasizes Text Material or Lecture Notes Some professors test almost exclusively on material covered in lecture. Others rely on their students’ learning most, or even all, of

the content in assigned chapters. Find out what your professor’s requirements are, because the way you study will vary accordingly. How to study when professors test lecture material If lectures are the main source of examination questions, make your lecture notes as complete and organized as possible. Before going to class, skim over the chapter, identifying key terms and examining the main figures, so that you can take effective lecture notes. Spend no more than 1 hour on this. Within 24 hours after class, rewrite (or type) your notes. Before rewriting, however, read the notes and make marginal notes about anything that is not clear. Then read the corresponding material in your text. Highlight or underline any sections that clarify questions you had in your notes. Read the entire chapter, including parts that are not covered in lecture. This extra information will give you breadth of understanding and will help you grasp key concepts. After reading the text, you are ready to rewrite your notes, incorporating relevant material from the text. It also helps to use the Glossary to find definitions for unfamiliar terms. Many students develop a set of flash cards of key terms and concepts as a way to study. Flash cards are a useful tool to help you learn scientific terminology. They are portable and can be used at times when other studying is not possible, for example, when riding a bus. Flash cards are not effective when the student tries to secondguess the professor. (“She won’t ask this, so I won’t make a flash card of it.”) Flash cards are also a hindrance when students rely on them exclusively. Studying flash cards instead of reading the text is a bit like reading the first page of each chapter in a mystery novel: it’s hard to fill in the missing parts, because you are learning the facts in a disconnected way. How to study when professors test material in the book If the assigned readings in the text are going to be tested, you must use your text intensively. After reading the chapter introduction, read the list of Learning Objectives for the first section. These objectives are written in behavioral terms; that is, they ask you to “do” something in order to demonstrate mastery. The objectives give you a concrete set of goals for each section of the chapter. At the end of each section, you will find Review questions keyed to the Learning Objectives. Test yourself, going back over the material to check your responses. Read each chapter section actively. Many students read and study passively. An active learner always has questions in mind and is constantly making connections. For example, there are many processes that must be understood in biology. Don’t try to blindly memorize these; instead, think about causes and effects so

that every process becomes a story. Eventually, you’ll see that many processes are connected by common elements. You will probably have to read each chapter two or three times before mastering the material. The second and third times through will be much easier than the first, because you’ll be reinforcing concepts that you have already partially learned. After reading the chapter, write a four- to six-page chapter outline by using the subheads as the body of the outline (first-level heads are boldface, in color, and all caps; second-level heads are in color and not all caps). Flesh out your outline by adding important concepts and boldface terms with definitions. Use this outline when preparing for the exam. Now it is time to test yourself. Answer the Test Your Understanding questions, and check your answers. Write answers to each of the Critical Thinking questions. Finally, review the Learning Objectives in the Chapter Summary, and try to answer them before reading the summary provided. If your professor has told you that some or all of the exam will be short-answer or essay format, write out the answer for each Learning Objective. Remember that this is a self-test. If you do not know an answer to a question, find it in the text. If you can’t find the answer, use the Index.

Learn the Vocabulary One stumbling block for many students is learning the many terms that make up the language of biology. In fact, it would be much more difficult to learn and communicate if we did not have this terminology, because words are really tools for thinking. Learning terminology generally becomes easier if you realize that most biological terms are modular. They consist of mostly Latin and Greek roots; once you learn many of these, you will have a good idea of the meaning of a new word even before it is defined. For this reason, we have included Appendix C, Understanding Biological Terms. To be sure you understand the precise definition of a term, use the Index and Glossary. The more you use biological terms in speech and writing, the more comfortable you will be.

Form a Study Group Active learning is facilitated if you do some of your studying in a small group. In a study group, the roles of teacher and learner can be interchanged: a good way to learn material is to teach. A study group lets you meet challenges in a nonthreatening environment and can provide some emotional support. Study groups are effective learning tools when combined with individual study of text and lecture notes. If, however, you and other members of your study group have not prepared for your meetings by studying individually in advance, the study session can be a waste of time.

Prepare for the Exam Your calendar tells you it is now 1 week before your first biology exam. If you have been following these suggestions, you are well prepared and will need only some last-minute reviewing. No allnighters will be required. During the week prior to the exam, spend 2 hours each day actively studying your lecture notes or chapter outlines. It helps many students to read these notes out loud (most people listen to what they say!). Begin with the first lecture/chapter covered on the exam, and continue in the order on the lecture syllabus. Stop when you have reached the end of your 2-hour study period. The following day, begin where you stopped the previous day. When you reach the end of your notes, start at the beginning and study them a second time. The material should be very familiar to you by the second or third time around. At this stage, use your textbook only to answer questions or clarify important points. The night before the exam, do a little light studying, eat a nutritious dinner, and get a full night’s sleep. That way, you’ll arrive in class on exam day with a well-rested body (and brain) and the self-confidence that goes with being well prepared. Eldra P. Solomon Linda R. Berg Diana W. Martin

A View of Life

NIBSC/Science Photo Library

1

KEY CONCEPTS

H1N1, the virus that causes H1N1 influenza (flu). H1N1 virus particles (blue) are visible on a cell (green). When this virus emerged, the human immune system was unfamiliar with its new combination of genes. As a result, the virus spread easily, causing a pandemic. The scanning electron micrograph (SEM) has been color-enhanced.

T

he H1N1 influenza (flu) outbreak became the focus of attention in April 2009. Within a few months, more than 200 countries around

1.1 Basic themes of biology include evolution, information transfer, and energy transfer.

the world had reported confirmed cases of this viral disease, and H1N1

1.2 Characteristics of life include cellular structure,

ease Control and Prevention (CDC), more than 200 known pathogens

growth and development, self-regulated metabolism, response to stimuli, and reproduction.

(disease-causing organisms) have the potential to strike globally. Histori-

1.3 Biological organization is hierarchical and includes chemical, cell, tissue, organ, organ system, and organism levels; ecological organization includes population, community, ecosystem, and biosphere levels.

had caused thousands of deaths. According to the U.S. Centers for Dis-

cally, new viral strains have claimed many human lives. For example, in 1918 an influenza pandemic killed more than 20 million people throughout the world. Epidemiologists warn that even today an influenza pandemic could kill millions of people. Pandemics such as H1N1 have negative

1.4 Information transfer includes DNA transfer of information from one generation to the next; chemical and electrical signals within and among the cells of every organism; and chemicals, visual displays, and sounds that allow organisms to communicate with one another and to interact with their environment.

global impact. They affect many aspects of life, including the global

1.5 Individual organisms and entire ecosystems depend on

lutionary relationships to known pathogens. For example, investigators

a continuous input of energy. Energy is transferred within cells and from one organism to another.

have determined that the 1918 flu pandemic was caused by an influenza A

1.6 Evolution is the process by which populations of organisms change over time, adapting to changes in their environment; the tree of life includes three major branches, or domains.

economy, travel, tourism, and education. Armed with new technology, biologists work closely with public health and other health-care professionals to prevent dangerous pandemics. When a new disease-causing agent emerges, biologists study its evo-

(H1N1) virus that may have mutated and newly emerged from a swine or avian host. The H1N1 strain that was identified in 2009 was related to the 1918 pathogen. Biologists determined that the 2009 H1N1 strain evolved from a com-

1.7 Biologists ask questions, develop hypotheses, make

bination of viruses that infected swine, birds, and humans. They found

predictions, and collect data by careful observation and experiment; based on their results, they come to conclusions.

that this strain of H1N1 contains unique combinations of gene segments for which humans do not have preexisting immunity. Knowledge about a

virus’s origins provides important clues to its structure and behavior, and suggests hypotheses for combating it. Scientists must then test their hypotheses in the laboratory. Researchers were able to determine the antigens (proteins) on the surface of H1N1. These antigens must combine with receptors on human cells in order to infect the cells. Based on careful studies of H1N1, a vaccine was quickly developed. Pathogens can strike quickly and spread rapidly, and the continuous evolution of drug-resistant pathogens presents a major challenge. New varieties of H1N1 continue to emerge, and investigators must quickly characterize them and assess their potential virulence. Scientists predict that new varieties may show increased drug resistance and may be more virulent. In addition, recently developed vaccines may no longer be effective. Emerging diseases will be discussed further in Chapter 24. This is an exciting time to study biology, the science of life. The remarkable new discoveries biologists are making almost daily affect every aspect of our lives, including our health, food, safety, relationships with humans and other organisms, and the environment of our planet. New knowledge provides new insights into the human species and the millions of other organisms with which we share this planet. Biology affects our personal, governmental, and societal decisions. For example, a combined effort at every level is necessary to provide the resources and knowledge to meet the

1. Evolution. Populations of organisms have evolved through time from earlier forms of life. Scientists have accumulated a wealth of evidence showing that the diverse life-forms on this planet are related and that populations have evolved, that is, have changed over time, from earlier forms of life. The process of evolution is the framework for the science of biology and is a major theme of this book. 2. Information transfer. Information must be transmitted within organisms and among organisms, and organisms must be able to receive information from their environment. The survival and function of every cell and every organism depend on the orderly transmission of information. Evolution depends on the transmission of genetic information from one generation to another. 3. Energy transfer. All life processes, including thousands of chemical transactions that maintain life’s organization, require a continuous input of energy. Most of the energy for life comes from sunlight. Energy from the sun is transferred through living systems from producers to consumers; decomposers obtain energy as they feed on the dead bodies and wastes of both producers and consumers. Energy is also continuously transferred from one chemical compound to another within every cell. Evolution, information transfer, and energy transfer are forces that give life its unique characteristics. We begin our study of biology by developing a more precise understanding of the fundamental characteristics of living systems.

challenges of global pandemics. Whatever your college major or career goals, knowledge of biological concepts is a vital tool for understanding our world and for

Review ■

meeting many of the personal, societal, and global challenges that confront us. Among these challenges are the expanding human

■

population, decreasing biological diversity, diminishing natural

Why are evolution, information transfer, and energy transfer considered basic to life? What does the term evolution mean as applied to populations of organisms?

resources, global climate change, and prevention and cure of diseases, such as heart disease, cancer, diabetes, Alzheimer’s disease, acquired immunodeficiency syndrome (AIDS), and influenza. Meet-

1.2 CHARACTERISTICS OF LIFE

ing these challenges will require the combined efforts of biologists

■ ■ LEARNING OBJECTIVE

and other scientists, health professionals, educators, politicians,

2

and biologically informed citizens. This book is a starting point for your exploration of biology. It will provide you with the basic knowledge and the tools to become a part of this fascinating science, as well as a more informed member of society.

1.1 THREE BASIC THEMES

Distinguish between living and nonliving things by describing the features that characterize living organisms.

We easily recognize that a pine tree, a butterfly, and a horse are living things, whereas a rock is not. Despite their diversity, the organisms that inhabit our planet share a common set of characteristics that distinguish them from nonliving things. These features include a precise kind of organization, growth and development, self-regulated metabolism, the ability to respond to stimuli, reproduction, and adaptation to environmental change.

■ ■ LEARNING OBJECTIVE

Organisms are composed of cells

1

Although they vary greatly in size and appearance, all organisms consist of basic units called cells. New cells are formed only by the division of previously existing cells. As will be discussed in Chapter 4, these concepts are expressed in the cell theory, a fundamental unifying concept of biology.

Describe three basic themes of biology.

In this first chapter we introduce three basic themes of biology. These themes are interconnected with one another and with almost every concept that we discuss in this book.

Mike Abbey/Visuals Unlimited, Inc.

Some of the simplest life-forms, such as protozoa, are unicellular organisms, meaning that each consists of a single cell (FIG. 1-1). In contrast, the body of a dog or a maple tree is made of billions of cells. In such complex multicellular organisms, life processes depend on the coordinated functions of component cells that may be organized to form tissues, organs, and organ systems. Every cell is enveloped by a protective plasma membrane that separates it from the surrounding external environment. The plasma membrane regulates passage of materials between the cell and its environment. Cells have specialized molecules that contain genetic instructions and transmit genetic information. In most cells, the genetic instructions are encoded in deoxyribonucleic acid, more simply known as DNA. Cells typically have internal structures called organelles that are specialized to perform specific functions. There are two fundamentally different types of cells: prokaryotic and eukaryotic. Prokaryotic cells are exclusive to bacteria and to microscopic organisms called archaea. All other organisms are characterized by their eukaryotic cells. These cells typically contain a variety of organelles enclosed by membranes, including a nucleus, which houses DNA. Prokaryotic cells are structurally simpler; they do not have a nucleus or other membrane-enclosed organelles.

250 μm

(a) Unicellular organisms consist of one intricate cell that performs all the functions essential to life. Ciliates, such as this Paramecium, move about by beating their hairlike cilia.

Biological growth involves an increase in the size of individual cells of an organism, in the number of cells, or in both. Growth may be uniform in the various parts of an organism, or it may be greater in some parts than in others, causing the body proportions to change as growth occurs. Some organisms—most trees, for example—continue to grow throughout their lives. Many animals have a defined growth period that terminates when a characteristic adult size is reached. An intriguing aspect of the growth process is that each part of the organism typically continues to function as it grows. Living organisms develop as well as grow. Development includes all the changes that take place during an organism’s life. Like many other organisms, every human begins life as a fertilized egg that then grows and develops. The structures and body form that develop are exquisitely adapted to the functions the organism must perform.

McMurray Photography

Organisms grow and develop

(b) Multicellular organisms, such as this African buffalo (Syncerus caffer) and the plants on which it grazes, may consist of billions of cells specialized to perform specific functions.

FIGURE 1-1 Unicellular and multicellular life-forms

Organisms regulate their metabolic processes Within all organisms, chemical reactions and energy transformations occur that are essential to nutrition, the growth and repair of cells, and the conversion of energy into usable forms. The sum of all the chemical activities of the organism is its metabolism. Metabolic processes occur continuously in every organism, and they must be carefully regulated to maintain homeostasis, an appropriate, balanced internal environment. When enough of a cell product has been made, its manufacture must be decreased or turned off. When a particular substance is required, cell processes that produce it must be turned on. These homeostatic mechanisms are self-regulating control systems that are remarkably sensitive and efficient.

The regulation of glucose (a simple sugar) concentration in the blood of complex animals is a good example of a homeostatic mechanism. Your cells require a constant supply of glucose molecules, which they break down to obtain energy. The circulatory system delivers glucose and other nutrients to all the cells. When the concentration of glucose in the blood rises above normal limits, glucose is stored in the liver and in muscle cells. When you have not eaten for a few hours, the glucose concentration begins to fall. Your body converts stored nutrients to glucose, bringing the glucose concentration in the blood back to normal levels. When the glucose concentration decreases, you also feel hungry and restore nutrients by eating.

Organisms respond to stimuli

A. B. Dowsett/Science Photo Library/Photo Researchers, Inc.

Flagella

1 μm

FIGURE 1-2 Biological movement

(a) Hairs on the leaf surface of the Venus flytrap (Dionaea muscipula) detect the touch of an insect, and the leaf responds by folding.

FIGURE 1-3 Plants respond to stimuli

David M. Dennis/Tom Stack & Associates

David M. Dennis/Tom Stack & Associates

These bacteria (Helicobacter pylori), equipped with flagella for locomotion, have been linked to stomach ulcers. The photograph was taken using a scanning electron microscope. The bacteria are not really red and blue. Their color has been artificially enhanced.

All forms of life respond to stimuli, physical or chemical changes in their internal or external environment. Stimuli that evoke a response in most organisms are changes in the color, intensity, or direction of light; changes in temperature, pressure, or sound; and changes in the chemical composition of the surrounding soil, air, or water. Responding to stimuli involves movement, though not always locomotion (moving from one place to another). In simple organisms, the entire individual may be sensitive to stimuli. Certain unicellular organisms, for example, respond to bright light by retreating. In some organisms, locomotion is achieved by the slow oozing of the cell, the process of amoeboid movement. Other organisms move by beating tiny, hairlike extensions of the cell called cilia or longer structures known as flagella (FIG. 1-2). Some bacteria move by rotating their flagella. Most animals move very obviously. They wiggle, crawl, swim, run, or fly by contracting muscles. Sponges, corals, and oysters have free-swimming larval stages, but most are sessile as adults, meaning that they do not move from place to place. In fact, they may remain firmly attached to a surface, such as the sea bottom or a rock. Many sessile organisms have cilia or flagella that beat rhythmically, bringing them food and oxygen in the surrounding water. Complex animals, such as grasshoppers, lizards, and humans, have highly specialized cells that respond to specific types of stimuli. For example, cells in the retina of the vertebrate eye respond to light. Although their responses may not be as obvious as those of animals, plants do respond to light, gravity, water, touch, and other stimuli. For example, plants orient their leaves to the sun and grow toward light. Many plant responses involve different growth rates of various parts of the plant body. A few plants, such as the Venus flytrap of the Carolina swamps, are very sensitive to touch and catch insects (FIG. 1-3). Their leaves are hinged along the midrib, and they have a scent that attracts insects. Trigger hairs on the leaf surface detect the arrival of an insect and stimulate the leaf to fold. When the edges come together, they interlock, preventing the insect’s escape. The leaf then secretes enzymes that kill and digest the insect. The Venus flytrap usually grows in nitrogendeficient soil. The plant obtains part of the nitrogen required for its growth from the insects it “eats.”

(b) The edges of the leaf come together and interlock, preventing the fly’s escape. The leaf then secretes enzymes that kill and digest the insect.

Organisms reproduce At one time, people thought worms arose spontaneously from horsehair in a water trough, maggots from decaying meat, and frogs from the mud

McMurray Photography

Cabisco/Visuals Unlimited, Inc.

100 μm

FIGURE 1-5 Adaptations These Burchell’s zebras (Equus burchelli), photographed in Tanzania, are behaviorally adapted to position themselves to watch for lions and other predators. Stripes are thought to be an adaptation for visual protection against predators. They serve as camouflage or to break up form when spotted from a distance. The zebra stomach is adapted for feeding on coarse grass passed over by other grazers, an adaptation that helps the animal survive when food is scarce.

(a) Asexual reproduction. One individual gives rise to two or more offspring that are similar to the parent. Difflugia, a unicellular amoeba, is shown dividing to form two amoebas.

L. E. Gilbert/Biological Photo Service

ther. This genetic variation is important in the vital processes of evolution and adaptation.

(b) Sexual reproduction. Typically, each of two parents contributes a gamete (sperm or egg). Gametes fuse to produce the offspring, which has a combination of the traits of both parents. A pair of tropical flies are shown mating.

FIGURE 1-4 Asexual and sexual reproduction

of the Nile. Thanks to the work of several scientists, including the Italian physician Francesco Redi in the 17th century and French chemist Louis Pasteur in the 19th century, we know that organisms arise only from previously existing organisms. Simple organisms, such as amoebas, perpetuate themselves by asexual reproduction (FIG. 1-4a). When an amoeba has grown to a certain size, it reproduces by splitting in half to form two new amoebas. Before an amoeba divides, its hereditary material (set of genes) is duplicated, and one complete set is distributed to each new cell. Except for size, each new amoeba is similar to the parent cell. The only way that variation occurs among asexually reproducing organisms is by genetic mutation, a permanent change in the genes. In most plants and animals, sexual reproduction is carried out by the fusion of an egg and a sperm cell to form a fertilized egg (FIG. 1-4b). The new organism develops from the fertilized egg. Offspring produced by sexual reproduction are the product of the interaction of various genes contributed by the mother and the fa-

Populations evolve and become adapted to the environment The ability of a population to evolve over many generations and adapt to its environment equips it to survive in a changing world. Adaptations are inherited characteristics that enhance an organism’s ability to survive in a particular environment. The long, flexible tongue of the frog is an adaptation for catching insects, the feathers and lightweight bones of birds are adaptations for flying, and the thick fur coat of the polar bear is an adaptation for surviving frigid temperatures. Adaptations may be structural, physiological, biochemical, behavioral, or a combination of all four (FIG. 1-5). Every biologically successful organism is a complex collection of coordinated adaptations produced through evolutionary processes.

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■

What characteristics distinguish a living organism from a nonliving object? What would be the consequences to an organism if its homeostatic mechanisms failed? Explain your answer. What do we mean by adaptations?

1.3 LEVELS OF BIOLOGICAL ORGANIZATION ■ ■ LEARNING OBJECTIVE 3

Construct a hierarchy of biological organization, including levels characteristic of individual organisms and levels characteristic of ecological systems.

Whether we study a single organism or the world of life as a whole, we can identify a hierarchy of biological organization (FIG. 1-6). At every level, structure and function are precisely coordinated. One way to study a particular level is by looking at its components. Biologists can gain insights about cells by studying atoms and molecules. Learning about a structure by studying its parts is called reductionism. However, the whole is more than the sum of its parts. Each level has emergent properties, characteristics not found at lower levels. Populations of organisms have emergent properties such as population density, age structure, and birth and death rates. The individuals that make up a population do not have these characteristics. Consider also the human brain. The brain is composed of millions of neurons (nerve cells). However, we could study all of these individual neurons and have no clue about the functional capacities of the brain. Only when the neurons are wired together in precise fashion are the emergent properties, such as the capacity for thought, judgment, and motor coordination, evident.

geographic area at the same time make up a population. The populations of various types of organisms that inhabit a particular area and interact with one another form a community. A community can consist of hundreds of different types of organisms. A community together with its nonliving environment is an ecosystem. An ecosystem can be as small as a pond (or even a puddle) or as vast as the Great Plains of North America or the Arctic tundra. All of Earth’s ecosystems together are known as the biosphere. The biosphere includes all of Earth that is inhabited by living organisms—the atmosphere, the hydrosphere (water in any form), and the lithosphere (Earth’s crust). The study of how organisms relate to one another and to their physical environment is called ecology (derived from the Greek oikos, meaning “house”).

Review ■ ■

What are the levels of organization within an organism? What are the levels of ecological organization?

Organisms have several levels of organization The chemical level, the most basic level of organization, includes atoms and molecules. An atom is the smallest unit of a chemical element that retains the characteristic properties of that element. For example, an atom of iron is the smallest possible amount of iron. Atoms combine chemically to form molecules. Two atoms of hydrogen combine with one atom of oxygen to form a single molecule of water. Although composed of two types of atoms that are gases, water can exist as a liquid or solid. The properties of water are very different from those of its hydrogen and oxygen components, an example of emergent properties. At the cellular level, many types of atoms and molecules associate with one another to form cells. However, a cell is much more than a heap of atoms and molecules. Its emergent properties make it the basic structural and functional unit of life, the simplest component of living matter that can carry on all the activities necessary for life. During the evolution of multicellular organisms, cells associated to form tissues. For example, most animals have muscle tissue and nervous tissue. Plants have epidermis, a tissue that serves as a protective covering, and vascular tissues that move materials throughout the plant body. In most complex organisms, tissues organize into functional structures called organs, such as the heart and stomach in animals and roots and leaves in plants. In animals, each major group of biological functions is performed by a coordinated group of tissues and organs called an organ system. The circulatory and digestive systems are examples of organ systems. Functioning together with great precision, organ systems make up a complex, multicellular organism. Again, emergent properties are evident. An organism is much more than its component organ systems.

Several levels of ecological organization can be identified Organisms interact to form still more complex levels of biological organization. All the members of one species living in the same

1.4 INFORMATION TRANSFER ■ ■ LEARNING OBJECTIVE 4 Summarize the importance of information transfer within and between living systems, giving specific examples.

An organism inherits the information it needs to grow, develop, carry on self-regulated metabolism, respond to stimuli, and reproduce. Each organism must also have precise instructions for making the molecules necessary for its cells to communicate. The information an organism requires to carry on these life processes is coded and transmitted in the form of chemical substances and electrical impulses. Organisms must also communicate information to one another.

DNA transmits information from one generation to the next Humans give birth only to human babies, not to giraffes or rosebushes. In organisms that reproduce sexually, each offspring is a combination of the traits of its parents. In 1953, James Watson and Francis Crick worked out the structure of DNA, the large molecule that makes up the genes, the units of hereditary material (FIG. 1-7). A DNA molecule consists of two chains of atoms twisted into a helix. As will be described in Chapter 2, each chain is made up of a sequence of chemical subunits called nucleotides. There are four types of nucleotides in DNA; and each sequence of three nucleotides is part of the genetic code. Watson and Crick’s work led to the understanding of this genetic code. The information coded in sequences of nucleotides in DNA transmits genetic information from generation to generation. The code works somewhat like an alphabet. The nucleotides can “spell” an amazing variety of instructions for making organisms as diverse as bacteria, frogs, and redwood trees. The genetic code is universal, that is, virtually identical in all organisms—a dramatic example of the unity of life.

Organism Organ systems work together in a functional organism.

Population A population consists of organisms of the same species.

Organism

Population

Organ system (e.g., skeletal system) Tissues and organs make up organ systems.

Organ system

Organ (e.g., bone) Tissues form organs.

Organ

Tissue (e.g., bone tissue) Cells associate to form tissues.

Community The populations of different species that populate the same area make up a community.

Community Tissue

Bone cells

Nucleus

Cell Cellular level Atoms and molecules make up the cytoplasm and form organelles, such as the nucleus and mitochondria (the site of many energy Organelle transformations). Organelles perform various functions of the cell.

Chemical level Atoms join to form molecules. Macromolecules are large molecules such as proteins and DNA.

Ecosystem A community together with the nonliving environment forms an ecosystem. Ecosystem

Macromolecule Biosphere Oxygen atom Molecule

Hydrogen atoms

Water

FIGURE 1-6 Animated The hierarchy of biological organization

Biosphere Earth and all of its communities constitute the biosphere.

Computer image of B-DNA by Geis/Stodola. Not to be reproduced without permission.

Cells use proteins and many other types of molecules to communicate with one another. In a multicellular organism, cells produce chemical compounds, such as hormones, that signal other cells. Hormones and other chemical messengers can signal cells in distant organs to secrete a particular required substance or change some metabolic activity. In this way chemical signals help regulate growth, development, and metabolic processes. The mechanisms involved in cell signaling often involve complex biochemical processes. Cell signaling is currently an area of intense research. A major focus has been the transfer of information among cells of the immune system. A better understanding of how cells communicate promises new insights into how the body protects itself against disease organisms. Learning to manipulate cell signaling may lead to new methods of delivering drugs into cells and new treatments for cancer and other diseases. Some organisms use electrical signals to transmit information. Most animals have nervous systems that transmit information by way of both electrical impulses and chemical compounds known as neurotransmitters. Information transmitted from one part of the body to another is important in regulating life processes. In complex animals, the nervous system gives the animal information about its outside environment by transmitting signals from sensory receptors such as the eyes and ears to the brain. Information must also be transmitted from one organism to another. Mechanisms for this type of communication include the release of chemicals, visual displays, and sounds. Typically, organisms use a combination of several types of communication signals. A dog may signal aggression by growling, using a particular facial expression, and laying its ears back. Many animals perform complex courtship rituals in which they display parts of their bodies, often elaborately decorated, to attract a mate.

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What is the function of DNA? What are two examples of cell signaling?

FIGURE 1-7 DNA DNA is the hereditary material that transmits information from one generation to the next. As shown in this model, DNA is a macromolecule that consists of two chains of atoms twisted into a helix. Each chain consists of subunits called nucleotides. The sequence of nucleotides makes up the genetic code.

1.5 THE ENERGY OF LIFE ■ ■ LEARNING OBJECTIVE 5

Information is transmitted by chemical and electrical signals Genes control the development and functioning of every organism. The DNA that makes up the genes contains the “recipes” for making all the proteins required by the organism. Proteins are large molecules important in determining the structure and function of cells and tissues. For example, brain cells differ from muscle cells in large part because they have different types of proteins. Some proteins are important in communication within and among cells. Certain proteins on the surface of a cell serve as markers so that other cells “recognize” them. Some cell-surface proteins serve as receptors that combine with chemical messengers.

Summarize the flow of energy through ecosystems and contrast the roles of producers, consumers, and decomposers.

Life depends on a continuous input of energy from the sun because every activity of a living cell or organism requires energy. Whenever energy is used to perform biological work, some is converted to heat and dispersed into the environment. Recall that all the energy transformations and chemical processes that occur within an organism are referred to as its metabolism. Energy is necessary to carry on the metabolic activities essential for growth, repair, and maintenance. Each cell of an organism requires nutrients that contain energy. During cellular respiration, cells capture energy released by nutrient molecules through a series of carefully regulated chemical reactions (FIG. 1-8). The cell can use this energy to do work, including the synthesis of

KEY POINT

Most of the energy for life is light energy from the sun, which is captured during photosynthesis; some of this energy is stored in the chemical bonds of glucose and other nutrients.

Light energy

Photosynthesis captures light energy

Oxygen

Energy stored in glucose and other nutrients

Carbon dioxide and water

Oxygen

Energy

Cellular respiration releases energy stored in glucose molecules

Synthesis of needed molecules and structures

Other Life Activities

r Homeostasis r Growth and development

r Reproduction r Movement of materials in and out of cells

r Movement of body FIGURE 1-8 Energy flow within and among organisms Algae and certain plant cells carry on photosynthesis, a process that uses light energy to produce glucose from carbon dioxide and water. Energy is stored in the chemical bonds of glucose and other nutrients produced from glucose. Through the process of cellular respiration, cells

required materials, such as new cell components. Virtually all cells carry on cellular respiration. Like individual organisms, ecosystems also depend on a continuous energy input. A self-sufficient ecosystem contains three types of organisms—producers, consumers, and decomposers— and includes a physical environment in which they can survive. These organisms depend on one another and on the environment

of all organisms, including algae and plant cells, then break down glucose and other nutrients. The energy released can be used to produce needed molecules and to fuel other life activities.

for nutrients, energy, oxygen, and carbon dioxide. There is a oneway flow of energy through ecosystems. Organisms can neither create energy nor use it with complete efficiency. During every energy transaction, some energy disperses into the environment as heat and is no longer available to the organism (FIG. 1-9). Plants, algae, and certain bacteria are producers, or autotrophs, that produce their own food from simple raw materials.

Light energy

Heat

Heat

Animals are consumers, or heterotrophs—that is, organisms that depend on producers for food, energy, and oxygen. Primary consumers eat producers, whereas secondary consumers eat primary consumers. Consumers obtain energy by breaking down sugars and other food molecules originally produced during photosynthesis. When chemical bonds are broken during this process of cellular respiration, their stored energy is made available for life processes: glucose + oxygen ¡ carbon dioxide + water + energy

Food

Primary consumer (caterpillar)

Heat

Heat Producer (plant)

Secondary consumer (robin)

Consumers contribute to the balance of the ecosystem. For example, consumers produce carbon dioxide required by producers. (Note that producers also carry on cellular respiration.) The metabolism of consumers and producers helps maintain the lifesustaining mixture of gases in the atmosphere. Most bacteria and fungi are decomposers, heterotrophs that obtain nutrients by breaking down nonliving organic material such as wastes, dead leaves and branches, and the bodies of dead organisms. In their process of obtaining energy, decomposers make the components of these materials available for reuse. If decomposers did not exist, nutrients would remain locked up in wastes and dead bodies, and the supply of elements required by living systems would soon be exhausted.

Review Plant litter, wastes

Soil

Dead bodies

Decomposers (bacteria, fungi)

FIGURE 1-9 Animated Energy flow through the biosphere Continuous energy input from the sun operates the biosphere. During photosynthesis, producers use the energy from sunlight to make complex molecules from carbon dioxide and water. Primary consumers, such as the caterpillar shown here, obtain energy, nutrients, and other required materials when they eat producers. Secondary consumers, such as the robin, obtain energy, nutrients, and other required materials when they eat primary consumers that have eaten producers. Decomposers obtain their energy and nutrients by breaking down wastes and dead organic material. During every energy transaction, some energy is lost to biological systems, dispersing into the environment as heat.

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1.6 EVOLUTION: THE BASIC UNIFYING CONCEPT OF BIOLOGY ■ ■ LEARNING OBJECTIVES 6 Demonstrate the binomial system of nomenclature by using specific

7 8 9

Most of these organisms use sunlight as an energy source and carry on photosynthesis, the process during which producers synthesize complex molecules such as glucose (a sugar) from carbon dioxide and water. The light energy is transformed into chemical energy, which is stored within the chemical bonds of the food molecules produced. Oxygen, which is required by the cells of most organisms including plant cells, is produced as a byproduct of photosynthesis: carbon dioxide + water + light energy ¡ glucose + oxygen

What components do you think a forest ecosystem might include? In what ways do consumers depend on producers? on decomposers? Include energy considerations in your answer.

examples and classify an organism (such as a human) in its domain, kingdom, phylum, class, order, family, genus, and species. Identify the three domains and the kingdoms of living organisms, and give examples of organisms assigned to each group. Give a brief overview of the scientific theory of evolution and explain why it is the principal unifying concept in biology. Apply the theory of natural selection to any given adaptation and suggest a logical explanation of how the adaptation may have evolved.

Evolution is the process by which populations of organisms change over time. Evolutionary theory has become the most important unifying concept of biology. As we will discuss, evolution involves passing genes for new traits from one generation to another, leading to differences in populations. The evolutionary perspective is important in every specialized field within biology. Biologists try to understand the structure, function, and behavior of organisms and their interactions with one another by considering them in light of the long, continuing process of evolution. Although we

discuss evolution in depth in Chapters 18 through 22, we present a brief overview here to give you the background necessary to understand other aspects of biology. First, we examine how biologists organize the millions of organisms that have evolved, and then we summarize some of the mechanisms that drive evolution.

Biologists use a binomial system for naming organisms Biologists have identified about 1.8 million species of extant (currently living) organisms and estimate that several million more remain to be discovered. To study life, we need a system for organizing, naming, and classifying its myriad forms. Systematics is the field of biology that studies the diversity of organisms and their evolutionary relationships. Taxonomy, a subspecialty of systematics, is the science of naming and classifying organisms. In the 18th century, Carolus Linnaeus, a Swedish botanist, developed a hierarchical system of naming and classifying organisms. Biologists still use this system today, with some modification. The species is a group of organisms with similar structure, function, and behavior. A species consists of one or more populations whose members are capable of breeding with one another; in nature, they do not breed with members of other species. Members of a population contribute to a common gene pool (all the genes present in the population) and share a common ancestry. Closely related species are grouped in the next broader category of classification, the genus (pl., genera). The Linnaean system of naming species is known as the binomial system of nomenclature because each species is assigned a two-part name. The first part of the name is the genus, and the second part, the specific epithet, designates a particular species belonging to that genus. The specific epithet is often a descriptive word expressing some quality of the organism. It is always used together with the full or abbreviated generic name preceding it. The generic name is always capitalized; the specific epithet is generally not capitalized. Both names are always italicized or underlined. For example, the domestic dog, Canis familiaris (abbreviated C. familiaris), and the timber wolf, Canis lupus (C. lupus), belong to the same genus. The domestic cat, Felis catus, belongs to a different genus. The scientific name of the American white oak is Quercus alba, whereas the name of the European white oak is Quercus robur. Another tree, the white willow, Salix alba, belongs to a different genus. The scientific name for our own species is Homo sapiens (“wise man”).

sists of 12 genera and about 34 living species. Family Canidae, along with family Ursidae (bears), family Felidae (catlike animals), and several other families that eat mainly meat, are all placed in order Carnivora. Order Carnivora, order Primates (to which chimpanzees and humans belong), and several other orders belong to class Mammalia (mammals). Class Mammalia is grouped with several other classes that include fishes, amphibians, reptiles, and birds in subphylum Vertebrata. The vertebrates belong to phylum Chordata, which is part of kingdom Animalia. Animals are assigned to domain Eukarya.

The tree of life includes three domains and several kingdoms Systematics has itself evolved as scientists have developed new techniques for inferring common ancestry among groups of organisms. As we will learn, biologists seek to classify organisms based on evolutionary relationships. These relationships are based on shared characteristics that distinguish a particular group. A group of organisms with a common ancestor is a clade. Systematists have developed a tree of life, a family tree showing proposed evolutionary relationships among organisms. These relationships are based on shared characteristics, including structural, developmental, behavioral, and molecular similarities, as well as on fossil evidence. FIGURE 1-11 is a cladogram, a branching diagram that depicts the tree of life as it is currently understood. As researchers report new findings, the classification of organisms changes and the branches of the tree of life must be redrawn. Although the tree of life is a work in progress, most biologists now assign organisms to three domains and to several kingdoms or clades. Bacteria have long been recognized as unicellular prokaryotic cells; they differ from all other organisms (except archaea) in that they are prokaryotes. Microbiologist Carl Woese (pronounced “woes”) has been a pioneer in developing molecular approaches to systematics. Woese and his colleagues selected a molecule known as small subunit ribosomal RNA (rRNA) that functions in the process of manufacturing proteins in all organisms. Because its molecular structure differs somewhat in various organisms, Woese hypothesized that the molecular composition of rRNA in closely related organisms would be more similar than in distantly related organisms. TABLE 1-1

Classification of the Cat, Human, and White Oak Tree

Taxonomic classification is hierarchical

Category

Cat

Human

White Oak

Just as closely related species may be grouped in a common genus, related genera can be grouped in a more inclusive group, a family. Families are grouped into orders, orders into classes, and classes into phyla (sing., phylum). Phyla can be assigned to kingdoms, and kingdoms are grouped in domains. Each formal grouping at any given level is a taxon (pl., taxa). Note that each taxon is more inclusive than the taxon below it. Together they form a hierarchy ranging from species to domain (TABLE 1-1 and FIG. 1-10). Consider a specific example. The family Canidae, which includes all doglike carnivores (animals that eat mainly meat), con-

Domain Kingdom Phylum Subphylum Class Order Family Genus Species

Eukarya Animalia Chordata Vertebrata Mammalia Carnivora Felidae Felis Felis catus

Eukarya Animalia Chordata Vertebrata Mammalia Primates Hominidae Homo Homo sapiens

Eukarya Plantae Anthophyta None Eudicotyledones Fagales Fagaceae Quercus Quercus alba

KEY POINT

Biologists use a hierarchical classification scheme with a series of taxonomic categories from species to domain; each category is more general and more inclusive than the one below it.

DOMAIN Eukarya

KINGDOM Animalia

PHYLUM Chordata

CLASS Mammalia

ORDER Primates

FAMILY Pongidae

GENUS Pan

SPECIES Pan troglodytes

FIGURE 1-10 Classification of the chimpanzee (Pan troglodytes)

This cladogram illustrates the evolutionary relationships among the three domains and among the major groups of organisms that belong to these domains.

5 μm

(a) The large, rodshaped bacterium Bacillus anthracis, a member of domain Bacteria, causes anthrax, a disease of cattle and sheep that can infect humans.

(b) These archaea (Methanosarcina mazei ), members of the domain Archaea, produce methane.

Archaea

Laurie Campbell/Getty Images

David M. Phillips/Visuals Unl.

CNRI/Science Photo Library/ Photo Researchers, Inc.

1 μm

Bacteria

Domain Eukarya

McMurray Photography

Domain Archaea R. Robinson/Visuals Unlimited, Inc.

Domain Bacteria

John Arnaldi

KEY POINT

10 μm

(c) These unicellular protozoa (Tetrahymena) are classified in one of the protist groups.

Protists

(d) Plants include many beautiful and diverse forms, such as the lady’s slipper (Phragmipedium caricinum).

(e) Among the fiercest animals, lions (Panthera leo) are also among the most sociable. The largest of the big cats, lions live in prides (groups).

Plants

Animals

(f) Mushrooms, such as these fly agaric mushrooms (Amanita muscaria), are fungi. The fly agaric is poisonous and causes delirium, raving, and profuse sweating when ingested. Fungi

Common ancestor of all organisms

FIGURE 1-11 Animated A survey of the three domains of life Biologists assign organisms to three domains and to several kingdoms and other groups. The protists do not form a clade and are no longer considered a kingdom. They are assigned to five “supergroups” (not shown).

Woese’s findings showed that there are two distinct groups of prokaryotes. He established the domain level of taxonomy and assigned the prokaryotes to two domains: Bacteria and Archaea (ar′-key-ah). The eukaryotes, organisms with eukaryotic cells, are classified in domain Eukarya. Woese’s work became widely accepted in the mid-1990s. In the classification system used in this book, each organism is assigned to a domain and to a kingdom or “supergroup.” Two

kingdoms correspond to the prokaryotic domains: kingdom Archaea corresponds to domain Archaea, and kingdom Bacteria corresponds to domain Bacteria. The remaining kingdoms and groups are assigned to domain Eukarya. Protists (for example, algae, slime molds, amoebas, and ciliates) are unicellular, colonial, or simple multicellular organisms that have a eukaryotic cell organization. The word protist, from the Greek for “the very first,” reflects the idea that protists were the first

eukaryotes to evolve. Protists are primarily aquatic organisms with diverse body forms, types of reproduction, modes of nutrition, and lifestyles. Some protists are adapted to carry out photosynthesis. Based mainly on molecular data that have clarified many evolutionary relationships among eukaryotes, the protists are no longer considered a kingdom. As we will learn in Chapter 26, several clades of protists have been identified, and many biologists now classify protists into five “supergroups.” Members of kingdom Plantae are complex multicellular organisms adapted to carry out photosynthesis. Among characteristic plant features are the cuticle (a waxy covering over aerial parts that reduces water loss) and stomata (tiny openings in stems and leaves for gas exchange); many plants have multicellular gametangia (organs that protect developing reproductive cells). Kingdom Plantae includes both nonvascular plants (mosses) and vascular plants (ferns, conifers, and flowering plants), those that have tissues specialized for transporting materials throughout the plant body. Most plants are adapted to terrestrial environments. Kingdom Fungi is composed of the yeasts, mildews, molds, and mushrooms. Fungi do not photosynthesize. They obtain their nutrients by secreting digestive enzymes into food and then absorbing the predigested food. Kingdom Animalia is made up of multicellular organisms that obtain their nutrition by eating other organisms. Most animals exhibit considerable cell and tissue specialization and body organization. These characters have evolved along with complex sense organs, nervous systems, and muscular systems. Most animals reproduce sexually; they have large, nonmotile (do not move from place to place) eggs and small sperm with flagella that propel them in their journey to find the egg. We have provided an introduction here to the groups of organisms that make up the tree of life. We will refer to them throughout this book, as we consider the many kinds of challenges organisms face and the various adaptations that have evolved in response to them. We discuss the diversity of life in more detail in Chapters 23 through 32, and we summarize classification in Appendix B.

ously existing forms. Darwin’s book raised a storm of controversy in both religion and science, some of which still lingers. Darwin’s theory of evolution has helped shape the biological sciences to the present day. His work generated a great wave of scientific observation and research that has provided much additional evidence that evolution is responsible for the great diversity of organisms on our planet. Even today, the details of evolutionary processes are a major focus of investigation and discussion. Darwin based his theory of natural selection on the following four observations: 1. Individual members of a species show some variation from one another. 2. Organisms produce many more offspring than will survive to reproduce (FIG. 1-12). 3. Because more individuals are produced than the environment can support, organisms must compete for necessary, but limited, resources such as food, sunlight, and space. Also, some organisms are killed by predators, disease organisms, or unfavorable natural conditions, such as weather changes. Which organisms are more likely to survive? 4. Individuals with characteristics that enable them to obtain and use resources, escape predators, resist disease organisms, and withstand changes in the environment are more likely to survive to reproductive maturity. The survivors that reproduce pass their adaptations for survival on to their offspring. Thus, the best-adapted individuals of a population leave, on average,

Species adapt in response to changes in their environment

J. Serrao/Photo Researchers, Inc.

Every organism is the product of numerous interactions between environmental conditions and the genes inherited from its ancestors. If all individuals of a species were exactly alike, any change in the environment might be disastrous to all, and the species would become extinct. Adaptations to changes in the environment occur as a result of evolutionary processes that take place over time and involve many generations.

Natural selection is an important mechanism by which evolution proceeds Although philosophers and naturalists discussed the concept of evolution for centuries, Charles Darwin and Alfred Wallace first brought a theory of evolution to general attention and suggested a plausible mechanism, natural selection, to explain it. In his book On the Origin of Species by Natural Selection, published in 1859, Darwin synthesized many new findings in geology and biology. He presented a wealth of evidence supporting his hypothesis that present forms of life descended, with modifications, from previ-

FIGURE 1-12 Egg masses of the wood frog (Rana sylvatica) Many more eggs are produced than can possibly develop into adult frogs. Random events are largely responsible for determining which of these developing frogs will hatch, reach adulthood, and reproduce. However, certain traits of each organism also contribute to the probability for success in its environment. Not all organisms are as prolific as the frog, but the generalization that more organisms are produced than survive is true throughout the living world.

(b) ‘I’iwi (Vestiaria cocciniea) in ‘ohi’a blossoms. The bill is adapted for feeding on nectar in tubular flowers.

Jack Jeffrey, Inc.

Jack Jeffrey, Inc.

Jack Jeffrey, Inc.

(a) The bill of this ‘Akiapola’au male (Hemignathus munroi ) is adapted for extracting insect larvae from bark. The lower mandible (jaw) is used to peck at and pull off bark, whereas the maxilla (upper jaw) and tongue remove the prey.

(c) Palila (Loxiodes bailleui ) in mamane tree. This finch-billed honeycreeper feeds on immature seeds in pods of the mamane tree. It also eats insects, berries, and young leaves.

FIGURE 1-13 Adaptation and diversification in Hawaiian honeycreepers All three species shown here are endangered, mainly because their habitats have been destroyed by humans or species introduced by humans.

more offspring than do other individuals. Because of this differential reproduction, a greater proportion of the population becomes adapted to the prevailing environmental conditions and challenges. The environment selects the best-adapted organisms for survival. Note that adaptation involves changes in populations rather than in individual organisms. Darwin did not know about DNA or understand the mechanisms of inheritance. Scientists now understand that most variations among individuals are a result of different varieties of genes that code for each characteristic. The ultimate source of these variations is random mutations, chemical or physical changes in DNA that persist and can be inherited. Mutations modify genes and by this process provide the raw material for evolution.

bills evolved (FIG. 1-13; see also Chapter 20 and Fig. 20-18). Some honeycreepers now have long, curved bills, adapted for feeding on nectar from tubular flowers. Others have short, thick bills for foraging for insects, and still others have adapted for eating seeds.

Review ■ ■

■

What is the binomial system of nomenclature? Biologists describe the tree of life as a work in progress. Why does the tree need to be modified? How might you explain the sharp claws and teeth of tigers in terms of natural selection?

1.7 THE PROCESS OF SCIENCE Populations evolve as a result of selective pressures from changes in their environment All the genes present in a population make up its gene pool. By virtue of its gene pool, a population is a reservoir of variation. Natural selection acts on individuals within a population. Selection favors individuals with genes specifying traits that allow them to respond effectively to pressures exerted by the environment. These organisms are most likely to survive and produce offspring. As successful organisms pass on their genetic recipe for survival, their traits become more widely distributed in the population. Over time, as populations continue to change (and as the environment itself changes, bringing different selective pressures), the members of the population become better adapted to their environment and less like their ancestors. As a population adapts to environmental pressures and exploits new opportunities for finding food, maintaining safety, and avoiding predators, the population diversifies and new species may evolve. The Hawaiian honeycreepers, a group of related birds, are a good example. When honeycreeper ancestors first reached Hawaii, few other birds were present, so there was little competition. Genetic variation among honeycreepers allowed some to move into different food zones, and over time, species with various types of

■ ■ LEARNING OBJECTIVES 10 Design a study to test a given hypothesis, using the procedure and terminology of the scientific method.

11 Compare the reductionist and systems approaches to biological research.

Biology is a science. The word science comes from a Latin word meaning “to know.” Science is a way of thinking and a method of investigating the natural world in a systematic manner. We test ideas, and based on our findings, we modify or reject these ideas. The process of science is investigative, dynamic, and often controversial. The observations made, the range of questions asked, and the design of experiments depend on the creativity of the individual scientist. Science, however, is influenced by cultural, social, historical, and technological contexts, so the process changes over time. The scientific method involves a series of ordered steps. Using the scientific method, scientists make careful observations, ask critical questions, and develop hypotheses, which are tentative explanations. Using their hypotheses, scientists make predictions that can be tested by making further observations or by performing experiments. They gather data, information that they can analyze, often using computers and sophisticated statistical methods.

They interpret the results of their experiments and draw conclusions from them. As we will discuss, scientists pose many hypotheses that cannot be tested by using all of the steps of the scientific method in a rigid way. Scientists use the scientific method as a generalized framework or guide. Biologists explore every imaginable aspect of life from the structure of viruses and bacteria to the interactions of the communities of our biosphere. Some biologists work mainly in laboratories, and others do their work in the field (FIG. 1-14). Perhaps you will decide to become a research biologist and help unravel the complexities of the human brain, discover new hormones that cause plants to flower, identify new species of animals or bacteria, or develop new stem cell strategies to treat cancer, AIDS, or heart disease. Applications of basic biological research have provided the technology to transplant kidneys, livers, and hearts; manipulate genes; treat many diseases; and increase world food production. Biology has been a powerful force in providing the quality of life that most of us enjoy. You may choose to enter an applied field of biology, such as environmental science, dentistry, medicine, pharmacology, or veterinary medicine. Many interesting careers in the biological sciences are discussed in the Careers section on our website.

Science requires systematic thought processes Science is systematic. Scientists organize and often quantify knowledge, making it readily accessible to all who wish to build on its foundation. In this way, science is both a personal and a social endeavor. Science is not mysterious. Anyone who understands

its rules and procedures can take on its challenges. What distinguishes science is its insistence on rigorous methods to examine a problem. Science seeks to give precise knowledge about the natural world; the supernatural is not accessible to scientific methods of inquiry. Science is not a replacement for philosophy, religion, or art. Being a scientist does not prevent one from participating in other fields of human endeavor, just as being an artist does not prevent one from practicing science. Deductive reasoning begins with general principles Scientists use two types of systematic thought processes: deduction and induction. With deductive reasoning, we begin with supplied information, called premises, and draw conclusions on the basis of that information. Deduction proceeds from general principles to specific conclusions. For example, if you accept the premise that all birds have wings and the second premise that sparrows are birds, you can conclude deductively that sparrows have wings. Deduction helps us discover relationships among known facts. A fact is information or knowledge based on evidence. Inductive reasoning begins with specific observations Inductive reasoning is the opposite of deduction. We begin with specific observations and draw a conclusion or discover a general principle. For example, you know that sparrows have wings, can fly, and are birds. You also know that robins, eagles, pigeons, and hawks have wings, can fly, and are birds. You might induce that all birds have wings and fly. In this way, you can use the inductive method to organize raw data into manageable categories by answering this question: What do all these facts have in common? A weakness of inductive reasoning is that conclusions generalize the facts to all possible examples. When we formulate the general principle, we go from many observed examples to all possible examples. This is known as an inductive leap. Without it, we could not arrive at generalizations. However, we must be sensitive to exceptions and to the possibility that the conclusion is not valid. For example, the kiwi bird of New Zealand does not have functional wings (FIG. 1-15). We can never conclusively prove a universal generalization. The generalizations in inductive conclusions come from the creative insight of the human mind, and creativity, however admirable, is not infallible.

Mark Moffett/Minden Pictures

Scientists make careful observations and ask critical questions

FIGURE 1-14 Biologist at work This biologist studying the rainforest canopy in Costa Rica is part of an international effort to study and preserve tropical rain forests. Researchers study the interactions of organisms and the effects of human activities on the rain forests.

In 1928, British bacteriologist Alexander Fleming observed that a blue mold had invaded one of his bacterial cultures. He almost discarded it, but then he noticed that the area contaminated by the mold was surrounded by a zone where bacterial colonies did not grow well. The bacteria were disease organisms of the genus Staphylococcus, which can cause boils and skin infections. Anything that could kill them was interesting! Fleming saved the mold, a variety of Penicillium (blue bread mold), and isolated the antibiotic penicillin from it. However, he had difficulty culturing it. Even though Fleming recognized the potential practical benefit of penicillin, he did not develop the chemical techniques needed to purify it, and more than ten years passed before the drug was

C. Dani/Peter Arnold

tive or negative. Test results should also be repeatable by independent observers. (3) It is falsifiable, which means it can be proven false, as we will discuss in the next section. After generating hypotheses, the scientist decides which, if any, could and should be subjected to experimental test. Why not test them all? Time and money are important considerations in conducting research. Scientists must establish priority among the hypotheses to decide which to test first.

FIGURE 1-15 Is this animal a bird? The kiwi bird of New Zealand is about the size of a chicken. Its tiny 2-inch wings cannot be used for flight. The survivor of an ancient order of birds, the kiwi has bristly, hairlike feathers and other characteristics that qualify it as a bird.

put to significant use. In 1939, Sir Howard Florey and Ernst Boris Chain developed chemical procedures to extract and produce the active agent penicillin from the mold. Florey took the process to laboratories in the United States, and penicillin was first produced to treat wounded soldiers in World War II. In recognition of their work, Fleming, Florey, and Chain shared the 1945 Nobel Prize in Physiology or Medicine.

Chance often plays a role in scientific discovery Fleming did not set out to discover penicillin. He benefited from the chance growth of a mold in one of his culture dishes. However, we may wonder how many times the same type of mold grew on the cultures of other bacteriologists who failed to make the connection and simply threw away their contaminated cultures. Fleming benefited from chance, but his mind was prepared to make observations and formulate critical questions, and his pen was prepared to publish them. Significant discoveries are usually made by those who are in the habit of looking critically at nature and recognizing a phenomenon or problem. Of course, the technology necessary for investigating the problem must also be available.

A falsifiable hypothesis can be tested In science, a well-stated hypothesis can be tested. If no evidence is found to support it, the hypothesis is rejected. The hypothesis can be shown to be false. Even results that do not support the hypothesis may be valuable and may lead to new hypotheses. If the results do support a hypothesis, a scientist may use them to generate related hypotheses. Let us consider a hypothesis that we can test by careful observation: female mammals (animals that have hair and produce milk for their young) bear live young. The hypothesis is based on the observations that dogs, cats, cows, lions, and humans all are mammals and all bear live young. Consider further that a new species, species X, is identified as a mammal based on its having hair and producing milk for its young. Biologists predict that females of species X will bear live young. (Is this inductive or deductive reasoning?) If a female of the new species gives birth to live offspring, the hypothesis is supported. Before the Southern Hemisphere was explored, most people would probably have accepted the hypothesis without question because all known furry, milk-producing animals did, in fact, bear live young. But biologists discovered that two Australian animals (the duck-billed platypus and the spiny anteater) had fur and produced milk for their young—but laid eggs (FIG. 1-16). The hypothesis, as stated, was false no matter how many times it had previously been supported. As a result, biologists either had to consider the

Scientists make careful observations, ask critical questions, and develop hypotheses. A hypothesis is a tentative explanation for observations or phenomena. Hypotheses can be posed as “if . . . then . . .” statements. For example, if students taking introductory biology attend classes, then they will make a higher grade on the exam than students who do not attend classes. In the early stages of an investigation, a scientist typically thinks of many possible hypotheses. A good hypothesis exhibits the following characteristics: (1) It is reasonably consistent with well-established facts. (2) It is capable of being tested; that is, it should generate definite predictions, whether the results are posi-

Tom McHugh/Photo Researchers, Inc.

A hypothesis is a testable statement

FIGURE 1-16 Is this animal a mammal? The duck-billed platypus (Ornithorhynchus anatinus) is classified as a mammal because it has fur and produces milk for its young. However, unlike most mammals, it lays eggs.

platypus and the spiny anteater as nonmammals or had to broaden their definition of mammals to include them. (They chose to do the latter.) A hypothesis is not true just because some of its predictions (the ones people happen to have thought of or have thus far been able to test) have been shown to be true. After all, they could be true by coincidence. In fact, a hypothesis can be supported by data, but it cannot really be proven true. An unfalsifiable hypothesis cannot be proven false; in fact, it cannot be scientifically investigated. Belief in an unfalsifiable hypothesis, such as the existence of invisible and undetectable elves, must be rationalized on grounds other than scientific ones.

that the nucleus is necessary for the metabolic processes that provide for growth and cell reproduction. But, the investigators asked, what if the operation itself, not the loss of the nucleus, caused the amoeba to die? They performed a controlled experiment, subjecting two groups of amoebas to the same operative trauma (FIG. 1-17). In the experimental group, the nucleus was removed; in the control group, it was not. Ideally, an experimental group differs from a control group only with respect to the variable being studied. In the control group, the researcher inserted a microloop into each amoeba and pushed it around inside the cell to simulate removal of the nucleus; then the instrument was withdrawn, leaving the nucleus inside. Amoebas treated with such a sham operation recovered and subsequently grew and divided. This experiment showed that the removal of the nucleus, not simply the operation, caused the death

Models are important in developing and testing hypotheses Hypotheses have many potential sources, including direct observations or even computer simulations. Increasingly in biology, hypotheses may KEY EXPERIMENT be derived from models that scientists have developed to provide a comprehenASK CRITICAL QUESTIONS: Why is the nucleus so large? What is its importance? sive explanation for a large number of DEVELOP HYPOTHESIS: Cells will be adversely affected if they lose their nuclei. observations. Examples of such testable models include the model of the strucPERFORM EXPERIMENTS: Using a microloop, researchers removed the nucleus from each ture of DNA and the model of the strucamoeba in the experimental group. Amoebas in the control group were subjected to the same surgical procedure, but their nuclei were not removed. ture of the plasma membrane (discussed in Chapter 5). The best design for an experiment can sometimes be established by performing computer simulations. Virtual testing and evaluation are undertaken before the experiment is performed in the laboratory or field. Modeling and computer simulation save time and money.

Many predictions can be tested by experiment A hypothesis is an abstract idea, but based on their hypotheses, scientists can make predictions that can be tested. For example, we might predict that biology students who study for ten hours will do better on an exam than students who do not study. As used here, a prediction is a deductive, logical consequence of a hypothesis. It does not have to be a future event. Some predictions can be tested by controlled experiments. Early biologists observed that the nucleus was the most prominent part of the cell, and they hypothesized that cells would be adversely affected if they lost their nuclei. Biologists predicted that if the nucleus were removed from the cell, then the cell would die. They then experimented by surgically removing the nucleus of a unicellular amoeba. The amoeba continued to live and move, but it did not grow, and after a few days it died. These results suggested

Amoeba dies

(a) Experimental group. When its nucleus is surgically removed with a microloop, the amoeba dies.

Amoeba lives

(b) Control group. A control amoeba subjected to similar surgical procedures (including insertion of a microloop), but without actual removal of the nucleus, does not die. RESULTS: Amoebas without nuclei died. Amoebas in the control group lived. CONCLUSION: Amoebas cannot live without their nuclei. The hypothesis is supported.

FIGURE 1-17 Testing a prediction regarding the importance of the nucleus Scientists observed that the nucleus was the most prominent part of the cell. They asked critical questions about their observation and developed the hypothesis that cells would be adversely affected if they lost their nuclei. Based on their hypothesis, they made a prediction that could be tested: if the nucleus is removed from an amoeba, the amoeba will die. The investigators then performed experiments to test this prediction. They studied their results and concluded that their hypothesis was supported.

of the amoebas. The conclusion is that amoebas cannot live without their nuclei. The results support the hypothesis that if cells lose their nuclei, they are adversely affected. We can conclude that the nucleus is essential for the survival of the cell.

Researchers must avoid bias In scientific studies, researchers must do their best to avoid bias or preconceived notions of what should happen. For example, to prevent bias, most medical experiments are carried out in a doubleblind fashion. When a drug is tested, one group of patients receives the new medication, and a control group of matched patients receives a placebo (a harmless starch pill similar in size, shape, color, and taste to the pill being tested). This is a double-blind study, named so because neither the patient nor the physician knows who is getting the experimental drug and who is getting the placebo. The pills or treatments are coded in some way, and the code is broken only after the experiment is over and the results are recorded. Not all experiments can be so neatly designed; for example, it is often difficult to establish appropriate controls.

Scientists interpret the results of experiments and make conclusions Scientists gather data in an experiment, interpret their results, and then draw conclusions from them. In the amoeba experiment described earlier, investigators concluded that the data supported the hypothesis that the nucleus is essential for the survival of the cell. When the results do support a hypothesis, scientists may use them to generate related hypotheses. Even results that do not support the hypothesis may be valuable and may lead to new hypotheses. Let us discuss another experiment. Research teams studying chimpanzee populations in Africa have reported that chimpanzees appear to learn specific ways to use tools from one another. Behavior that is learned from others in a population and passed to future generations is what we call “culture.” In the past, most biologists have thought that only humans had culture. It has been difficult to test this type of learning in the field, and the idea has been controversial. Biologists have asked critical questions about whether chimpanzees learned how to use tools by observing one another. Investigators at Yerkes National Primate Research Center in Atlanta developed a hypothesis that chimpanzees can learn particular ways to use tools by observing other chimps. They predicted that if they taught one chimp to use a stick to obtain food from a dispenser, other chimps would learn the technique from the educated one (FIG. 1-18). These researchers divided chimpanzees into two experimental groups with 16 in each group. Then they taught a highranking female in each group to use a stick to obtain food from an apparatus. The two chimps were taught different methods. One chimp was taught to poke the stick inside the device to free the food. The other was taught to use the stick to lift a hook that removed a blockage, allowing the food to roll forward out of the device. A third group served as a control group. The chimps in the control group were given access to the sticks and the apparatus with the food inside but none were taught how to use the sticks. All of the control-group chimps manipulated the apparatus with the stick, but none succeeded in releasing food.

When the chimps were returned to their groups, other chimps observed how the educated chimps used the stick, and a large majority began to use sticks in the same way. The chimps in each group learned the specific style of using the stick that their educated chimp had been taught. Most used the stick to obtain food at least 10 times. Two months later, the apparatus was reintroduced to the chimps. Again, most of the chimps used the learned technique for obtaining food. The results of the experiment supported the hypothesis. The researchers concluded that chimpanzees are capable of culturally transmitting learned technology. Sampling error can lead to inaccurate conclusions One reason for inaccurate conclusions is sampling error. Because not all cases of what is being studied can be observed or tested (scientists cannot study every amoeba or every chimpanzee population), scientists must be content with a sample. How can scientists know whether that sample is truly representative of whatever they are studying? If the sample is too small, it may not be representative because of random factors. A study with only two, or even nine, amoebas may not yield reliable data that can be generalized to other amoebas. If researchers test a large number of subjects, they are more likely to draw accurate scientific conclusions (FIG. 1-19). The scientist seeks to state with some level of confidence that any specific conclusion has a certain statistical probability of being correct. Experiments must be repeatable When researchers publish their findings in a scientific journal, they typically describe their methods and procedures in sufficient detail so that other scientists can repeat the experiments. When the findings are replicated, the conclusions are, of course, strengthened.

A theory is supported by tested hypotheses Nonscientists often use the word theory incorrectly to refer to a hypothesis or even to some untestable idea they wish to promote. A scientific theory is actually an integrated explanation of some aspect of the natural world that is based on a number of hypotheses, each supported by consistent results from many observations or experiments. A scientific theory relates data that previously appeared unrelated. A good theory grows, building on additional facts as they become known. It predicts new facts and suggests new relationships among phenomena. It may even suggest practical applications. A scientific theory, by showing the relationships among classes of facts, simplifies and clarifies our understanding of the natural world. As Einstein wrote, “In the whole history of science from Greek philosophy to modern physics, there have been constant attempts to reduce the apparent complexity of natural phenomena to simple, fundamental ideas and relations.” Developing scientific theories is indeed a major goal of science.

Many hypotheses cannot be tested by direct experiment Some well-accepted theories do not lend themselves to hypothesis testing by ordinary experiments. Often, these theories describe events that occurred in the distant past. We cannot directly observe

KEY EXPERIMENT ASK CRITICAL QUESTIONS: Do chimpanzees learn how to use tools by observing one another? DEVELOP HYPOTHESIS: Chimpanzees can learn particular ways to use tools by observing other chimps. PERFORM EXPERIMENTS: One female in each of two groups of 16 chimps was educated in a specific way to use a stick to obtain food. The two educated chimps were then returned to their respective groups. Chimpanzees in a control group were not taught how to use a stick.

Number of chimps

12

8

4

0

Control

Exp. 1

Exp. 2

(a) Number of chimpanzees successfully employing specific method of tool use.

RESULTS: Chimpanzees in each experimental group observed the use of the stick by the educated chimp, and a majority began to use the stick in the same way. When tested two months later, many of the chimps in each group continued to use the stick. The results presented here have been simplified and are based on the number of chimps observed to use the learned method at least 10 times. All but one chimp in each group learned the technology, but a few used it only a few times. Some chimps taught themselves the alternative method and used that alternative. However, most conformed to the group’s use of the method that the investigator taught to the educated chimp. CONCLUSION: Chimpanzees learn specific ways to use tools by observing other chimps. The hypothesis was supported.

the origin of the universe from a very hot, dense state about 13.7 billion years ago (the Big Bang theory). However, physicists and cosmologists have been able to formulate many hypotheses related to the Big Bang and to test many of the predictions derived from these hypotheses. Similarly, humans did not observe the evolution of major groups of organisms because that process took place over millions of years and occurred before humans had evolved. However, many hypotheses about evolution have been posed, and predictions

Number of chimps

David Bygott

12

8

4

0

Control

Exp. 1

Exp. 2

(b) Number of chimpanzees successfully employing learned method of tool use two months later.

FIGURE 1-18 Testing a prediction about learning in chimpanzee populations In the photo, wild chimpanzees are shown observing a member of their group using a tool.

based on them have been tested. For example, if complex organisms evolved from simple life-forms, we would find the fossils of the simplest organisms in the oldest strata (rock layers). As we explore more recent strata, we would expect to find increasingly complex organisms. Indeed, scientists have found this progression of simple to complex fossils. In addition to fossils, evidence for evolution comes from many sources, including physical and molecular similarities between organisms. Evidence also comes from recent and current studies

Curtain

forms—bacteria and protists—that were neither plant nor animal. Biologists had to make a paradigm shift, that is, they changed their view of reality, to accommodate this new knowledge. They assigned these newly discovered organisms to new kingdoms. In a more recent paradigm shift, biologists have revised their idea that we are born with all of the brain cells we will ever have. We now understand that certain areas of the brain continue to produce new neurons throughout life.

Single selection

Marbles

Systems biology integrates different levels of information

produces

Assumption

100% blue

Actual ratio 20% blue 80% white

(a) Taking a single selection can result in sampling error. If the only marble selected is blue, we might assume all the marbles are blue. Curtain

Marbles

Multiple selections produce

Assumption

Actual ratio 20% blue 80% white

30% blue 70% white

(b) The greater the number of selections we take of an unknown, the more likely we can make valid assumptions about it.

FIGURE 1-19 Animated Statistical probability

of evolution in action. Many aspects of ongoing evolution can, in fact, be studied in the laboratory or in the field. The evidence for evolution is so compelling that virtually all scientists today accept evolutionary theory as an integral part of biology.

Paradigm shifts allow new discoveries A paradigm is a set of assumptions or concepts that constitute a way of thinking about reality. For example, from the time of Aristotle to the mid-19th century, biologists thought that organisms were either plants (kingdom Plantae) or animals (kingdom Animalia). This concept was deeply entrenched. However, with the development of microscopes, investigators discovered tiny life-

In the reductionist approach to biology, researchers study the simplest components of biological processes. Their goal is to synthesize their knowledge of many small parts to understand the whole. Reductionism has been (and continues to be) important in biological research. However, as biologists and their tools have become increasingly sophisticated, huge amounts of data have been generated, bringing the science of biology to a different level. Systems biology is a field of biology that builds on information provided by the reductionist approach and develops large data sets, typically generated by computers. Systems biology is also referred to as integrative biology. Reductionism and systems biology are complementary approaches. Using reductionism, biologists have discovered basic information about components, such as molecules, genes, cells, and organs. Systems biologists, who focus on systems as a whole rather than on individual components, need this basic knowledge to study, for example, the interactions among various parts and levels of an organism. Systems biologists integrate data from various levels of complexity with the goal of understanding the big picture—how biological systems function. For example, systems biologists are developing models of different aspects of cell function. Normal cell function depends on the precise actions of hundreds of proteins that relay signals received from other cells. Proteins also relay signals from one part of the cell to another. Researchers are producing detailed maps of the molecular pathways that maintain cell function (FIG. 1-20). One group of researchers has developed a model consisting of nearly 8000 chemical signals involved in a molecular network that leads to programmed cell death. By understanding cell communication, the interactions of genes and proteins in metabolic pathways, and physiological processes, systems biologists hope to eventually develop a model of the whole organism. Systems biology is increasingly used to study disease processes. For example, the interactions between the pathogen and the host cell can be mapped. The development of systems biology was fueled by the huge amount of data generated by the Human Genome Project. Researchers working on this project identified the DNA sequences that make up the estimated 25,000 genes of the human genome, the complete set of genes that make up the human genetic material. Computer software developed for the Human Genome Project can analyze very large data sets. These programs are being used to integrate data about protein interactions and many other aspects of molecular biology. Systems biologists view biology in terms of information systems. Increasingly, they depend on mathematics, statistics, and engineering principles.

Hawoong Jeong, University of Notre Dame/Science Photo Library

FIGURE 1-20 Map showing protein interactions in a yeast cell (Saccharomyces cerevisiae) Each dot represents one protein in this single-celled fungus. Lines connect proteins that interact with one another. Highly connected proteins are more critical to cell function. Many of these interactions have also been identified in human cells. Specific proteins were removed by removing the genes that coded for their production. The proteins have been color-coded according to the effect their removal has on the yeast cell: red, lethal; green, not lethal; orange, slower growth; yellow, unknown effects.

Science has ethical dimensions Scientific investigation depends on a commitment to practical ideals, such as truthfulness and the obligation to communicate results. Honesty is particularly important in science. Consider the great (though temporary) damage done whenever an unprincipled or even desperate researcher, whose career may depend on the publication of a research study, knowingly disseminates false data. Until the deception is uncovered, researchers may devote thousands of dollars and hours of precious professional labor to futile lines of research inspired by erroneous reports. Deception can also be dangerous, especially in medical research. Fortunately, science tends to correct itself through consistent use of the scientific process. Sooner or later, someone’s experimental results are sure to cast doubt on false data. Science and technology continuously interact. As scientists doing basic research report new findings, engineers and other in-

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ventors develop new products. Many of those products contribute to our quality of life. New technology also provides scientists with more powerful tools for their research and increases the potential for new discoveries. For example, a few years ago determining the genome of any eukaryote required several rooms filled with machines that could sequence the genes. This endeavor also cost millions of dollars. New technologies have revolutionized gene sequencing, allowing complex genomes to be determined quickly, with less equipment, and at far less expense. Plans are under way to determine the genomes of 10,000 vertebrates. Science and technology continue to change society, and these changes present new challenges. In addition to being ethical about their own work, scientists face many societal and political issues surrounding areas such as genetic research, stem cell research, cloning, and human and animal experimentation. For example, some stem cells that show great potential for treating human disease come from early embryos. The cells can be taken from fiveor six-day-old human embryos and then cultured in laboratory glassware. At this stage, the embryo is a group of cells about 0.15 mm long (0.006 in). Such cells could be engineered to treat failing hearts or brains harmed by stroke, injury, Parkinson’s disease, or Alzheimer’s disease. They could save the lives of burn victims and perhaps be engineered to treat specific cancers. Scientists, and the larger society, will need to determine whether the potential benefits of any type of research outweigh its ethical risks. The era of the genome brings with it many ethical concerns and responsibilities. How do people safeguard the privacy of genetic information? For example, suppose that you have a family history of breast cancer and learn from genetic testing that you have one of the BRCA mutations. These mutations increase risk for developing breast cancer and certain other cancers. How can you be certain that knowledge of your individual genetic code would not be used against you when you seek employment or health insurance? Scientists must be ethically responsible and must help educate people about their work, including its benefits relative to its risks. It is significant that at the very beginning of the Human Genome Project, part of its budget was allocated for research on the ethical, legal, and social implications of its findings.

Review ■ ■ ■

What are the characteristics of a good hypothesis? What is meant by a “controlled” experiment? What is systems biology?

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

1.1 (page 2) 1 Describe three basic themes of biology. ■

Three basic themes of biology are evolution, transfer of information, and energy transfer. The process of evolution results in populations changing over time and explains how the ancestry of organisms can be traced back to earlier forms of life. Information must be transmitted within cells, among cells, among organisms, and from one generation to the next. Life requires continuous input of energy from the sun.

1.2 (page 2) 2 Distinguish between living and nonliving things by describing the features that characterize living organisms. ■ Every living organism is composed of one or more cells. Living things grow by increasing the size and/or number of their cells. Metabolism includes all the chemical activities that take place ■ in the organism, including the chemical reactions essential to nutrition, growth and repair, and conversion of energy to usable forms. Homeostasis refers to the appropriate, balanced internal environment.

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Organisms respond to stimuli, physical or chemical changes in their external or internal environment. Responses typically involve movement. In asexual reproduction, offspring are typically identical to the single parent, except for size; in sexual reproduction, offspring are the product of the fusion of gametes, and genes are typically contributed by two parents. As populations evolve, they become adapted to their environment. Adaptations are traits that increase an organism’s ability to survive in its environment.

1.3 (page 5) 3 Construct a hierarchy of biological organization, including levels characteristic of individual organisms and levels characteristic of ecological systems. ■ Biological organization is hierarchical. A complex organism is organized at the chemical, cell, tissue, organ, organ system, and organism levels. Cells associate to form tissues that carry out specific functions. In most multicellular organisms, tissues organize to form functional structures called organs. An organized group of tissues and organs form an organ system. Functioning together, organ systems make up a complex, multicellular organism.

1.5 (page 8) 5 Summarize the flow of energy through ecosystems and contrast the roles of producers, consumers, and decomposers. ■ Activities of living cells require energy. Life depends on continuous energy input from the sun. During photosynthesis, plants, algae, and certain bacteria use the energy of sunlight to synthesize complex molecules from carbon dioxide and water. ■ Virtually all cells carry on cellular respiration, a biochemical process in which they capture the energy stored in nutrients by producers. Some of that energy is then used to synthesize required materials or to carry out other cell activities. ■ A self-sufficient ecosystem includes producers, or autotrophs, which make their own food; primary consumers, which eat producers, and typically secondary consumers that eat primary consumers; and decomposers, which obtain energy by breaking down wastes and dead organisms. Consumers and decomposers are heterotrophs, organisms that depend on producers as an energy source and for food and oxygen. Learn more about energy flow by clicking on the figure in CengageNOW.

1.6 (page 10) 6 Demonstrate the binomial system of nomenclature by using specific

Organ

Bone cells Tissue

Nucleus

Cell

Organelle

■

The basic unit of ecological organization is the population. Various populations form communities; a community and its physical environment are an ecosystem; all of Earth’s ecosystems together make up the biosphere. Learn more about biological organization by clicking on the figure in CengageNOW.

1.4 (page 7) 4 Summarize the importance of information transfer within and between living systems, giving specific examples. Organisms transmit information chemically, electrically, and behaviorally. ■ DNA, which makes up the genes, is the hereditary material. Information encoded in DNA is transmitted from one generation to the next. DNA contains the instructions for the development of an organism and for carrying out life processes. DNA codes for proteins, which are important in determining the structure and function of cells and tissues. ■ Hormones, chemical messengers that transmit messages from one part of an organism to another, are important in cell signaling. ■ Many organisms use electrical signals to transmit information; most animals have nervous systems that transmit electrical impulses and release neurotransmitters. ■

examples and classify an organism (such as a human) in its domain, kingdom, phylum, class, order, family, genus, and species. ■ Millions of species have evolved. A species is a group of organisms with similar structure, function, and behavior that, in nature, breed only with one another. Members of a species contribute to a common gene pool and share a common ancestry. ■ Biologists use a binomial system of nomenclature in which the name of each species includes a genus name and a specific epithet. Taxonomic classification is hierarchical; it includes species, genus, family, order, class, phylum, kingdom, and domain. Each grouping is referred to as a taxon. A group of organisms with a common ancestor is a clade. 7 Identify the three domains and the kingdoms of living organisms, and give examples of organisms assigned to each group. ■ Bacteria and archaea have prokaryotic cells; all other organisms have eukaryotic cells. Prokaryotes make up two of the three domains. ■ Organisms are classified in three domains: Archaea, Bacteria, and Eukarya and several kingdoms: Archaea, Bacteria, Fungi (e.g., molds and yeasts), Plantae, and Animalia. Protists (e.g., algae, water molds, slime molds, and amoebas) are now assigned to five “supergroups.” Learn more about life’s diversity by clicking on the figure in CengageNOW.

8 Give a brief overview of the scientific theory of evolution and explain why it is the principal unifying concept in biology. Evolution is the process by which populations change over time in ■ response to changes in the environment. Evolutionary theory explains how millions of species came to be and helps us understand the structure, function, behavior, and relationships of organisms. ■ Natural selection, the major mechanism by which evolution proceeds, favors individuals with traits that enable them to cope with environmental changes. Charles Darwin based his theory of natural selection on his observations that individuals of a species vary; organisms produce more offspring than survive to reproduce; organisms must compete for limited resources; and individuals that are best adapted to their environment are more likely to survive and reproduce, and pass on their hereditary information. Their traits become more widely distributed in the population. ■ The source of variation in a population is random mutation.

9 Apply the theory of natural selection to any given adaptation and suggest a logical explanation of how the adaptation may have evolved. ■ When the ancestors of Hawaiian honeycreepers first reached Hawaii, few other birds were present, so there was little competition for food. Through many generations, honeycreepers with longer, more curved bills became adapted for feeding on nectar from tubular flowers. Perhaps those with the longest, most curved bills were best able to survive in this food zone and lived to transmit their genes to their offspring. Those with shorter, thicker bills were more successful foraging for insects and passed their genes to new generations of offspring. Eventually, different species evolved that were adapted to specific food zones.

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1.7 (page 15)

Do your own random sampling by clicking on the figure in CengageNOW.

10 Design a study to test a given hypothesis, using the procedure and terminology of the scientific method. The process of science is a dynamic approach to investigation. The scientific method is a general framework that scientists use in their work; it includes observing, recognizing a problem or stating a critical question, developing a hypothesis, making a prediction that can be tested, making further observations, performing experiments, interpreting results, and drawing conclusions that support or falsify the hypothesis. ■ Deductive reasoning and inductive reasoning are two categories of systematic thought used in the scientific method. Deductive reasoning proceeds from general principles to specific conclusions ■

and helps us discover relationships among known facts. Inductive reasoning begins with specific observations and draws conclusions from them. Inductive reasoning helps us discover general principles. A hypothesis is a tentative explanation for observations or phenomena. A hypothesis can be tested. If no evidence is found to support it, the hypothesis is rejected. A well-designed scientific experiment typically includes both a control group and an experimental group and must be as free as possible from bias. The control group should be as closely matched to the experimental group as possible. Ideally, the experimental group differs from the control group only with respect to the variable being studied.

A scientific theory is an integrated explanation of some aspect of the natural world that is based on a number of hypotheses, each supported by consistent results from many observations or experiments. 11 Compare the reductionist and systems approaches to biological research. ■ Using reductionism, researchers study the simplest components of biological processes, for example, molecules or cells. Systems biology uses knowledge provided by reductionism. Systems biologists integrate data from various levels of complexity with the goal of understanding how biological systems function. ■

T E S T YO U R U N D E R S TA N D I N G 1. Homeostasis (a) is the appropriate, balanced internal environment (b) is the sum of all the chemical activities of an organism (c) is the long-term response of organisms to changes in their environment (d) occurs at the ecosystem level, not in cells or organisms (e) may be sexual or asexual

7. Cellular respiration (a) is a process whereby sunlight is used to synthesize cell components with the release of energy (b) occurs in heterotrophs only (c) is carried on by both autotrophs and heterotrophs (d) causes chemical changes in DNA (e) occurs in response to environmental changes

2. An amoeba splits into two smaller amoebas. This is an example of (a) homeostasis (b) neurotransmission (c) asexual reproduction (d) sexual reproduction (e) metabolism

8. Fungi are assigned to domain (a) Protista (b) Archaea (c) Bacteria (d) Eukarya (e) Plantae

3. Cells (a) are the building blocks of living organisms (b) always have nuclei (c) are not found among the bacteria (d) a, b, and c (e) a and b 4. Which of the following is a correct sequence of levels of biological organization? 1. organ system 2. chemical 3. tissue 4. organ 5. cell (a) 2, 3, 5, 4, 1 (b) 5, 3, 4, 1, 2 (c) 2, 5, 3, 1, 4 (d) 2, 5, 3, 4, 1 (e) 5, 2, 3, 4, 1 5. Which of the following is a correct sequence of levels of ecological organization? 1. community 2. organism 3. ecosystem 4. population 5. biosphere (a) 5, 3, 1, 4, 2 (b) 2, 4, 1, 3, 5 (c) 2, 1, 4, 3, 5 (d) 4, 2, 1, 3, 5 (e) 2, 4, 3, 1, 5 6. DNA (a) is produced during cellular respiration (b) functions mainly to transmit information from one species to another (c) cannot be changed (d) is a neurotransmitter (e) makes up the genes

9. The scientific name for corn is Zea mays. Zea is the (a) specific epithet (b) genus (c) class (d) kingdom (e) phylum 10. Darwin suggested that evolution takes place by (a) mutation (b) changes in the individuals of a species (c) natural selection (d) interaction of hormones during competition for resources (e) homeostatic responses to each change in the environment 11. Ideally, an experimental group differs from a control group (a) only with respect to the hypothesis being tested (b) because its subjects are more reliable (c) in that it is less subject to bias (d) in that it is less vulnerable to sampling error (e) only with respect to the variable being studied 12. A systems biologist would most likely work on (a) better understanding the components of cells (b) developing a better system of classification of organisms (c) devising a new series of steps for the scientific method (d) researching a series of reactions that communicate information in the cell (e) identifying the connections and interactions of neurons in order to learn about brain function

13. Identify the domain and kingdoms by writing the name for each on the blank lines. Bacteria

Archaea Protist groups

Common ancestor of all organisms

CRITICAL THINKING 1. What would happen if a homeostatic mechanism failed? Describe a homeostatic mechanism at work in your body (other than the regulation of glucose cited in the chapter). 2. Contrast the reductionist approach with systems biology. How are the two approaches complementary? Which approach is more likely to consider emergent properties? 3. What are some characteristics of a good hypothesis? Give an example. 4. Make a prediction and devise a suitably controlled experiment to test each of the following hypotheses: (a) A type of mold found in your garden produces an effective antibiotic. (b) The growth rate of a bean seedling is affected by temperature. (c) Estrogen alleviates symptoms of Alzheimer’s disease in elderly women. 5. EVOLUTION LINK. In what ways does evolution depend on transfer of information? In what ways does transfer of information depend on evolution? 6. EVOLUTION LINK. How might an understanding of evolutionary processes help a biologist doing research in (a) animal

behavior, for example, the hunting behavior of lions? (b) the development of a new antibiotic to replace one to which bacteria have become resistant? (c) conservation of a specific plant in a rain forest? 7. ANALYZING DATA. Compare the two graphs in Figure 1-18. What information does the second graph illustrate? What possible explanation can you give for the differences shown in the two graphs? 8. SCIENCE, TECHNOLOGY, AND SOCIETY. New technology is allowing scientists to develop effective gene therapies to treat serious diseases. In the future gene therapy may be used to treat such conditions as addiction or bipolar disorder. What if gene therapy could be used to produce children with athletic ability, artistic ability, or high IQ? Do you have any ethical concerns about these possibilities? If so, where and how would you draw the line? Additional questions are available in CengageNOW at www.cengage.com/ login.

2

Atoms and Molecules: The Chemical Basis of Life

Frans Lanting/Minden Pictures

Water is a basic requirement for all life. A jaguar (Panthera onca), the largest cat in the Western Hemisphere, pauses to drink water from a rainforest stream.

KEY CONCEPTS

A

2.1 Carbon, hydrogen, oxygen, and nitrogen are the most

forest, as well as abundant unseen insects and microorganisms, share

abundant elements in living things.

fundamental similarities in their chemical composition and basic meta-

2.2 The chemical properties of an atom are determined by

bolic processes. These chemical similarities provide strong evidence for

its highest-energy electrons, known as valence electrons.

the evolution of all organisms from a common ancestor and explain why

2.3 A molecule consists of atoms joined by covalent

much of what biologists learn from studying bacteria or rats in laborato-

bonds. Other important chemical bonds include ionic bonds. Hydrogen bonds and van der Waals interactions are weak attractions.

2.4 The energy of an electron is transferred in a redox reaction.

knowledge of chemistry is essential for understanding organisms and how they function. This jaguar and the plants of the tropical rain

ries can be applied to other organisms, including humans. Furthermore, the basic chemical and physical principles governing organisms are not unique to living things, for they apply to nonliving systems as well. The success of the Human Genome Project and related studies has

2.5 Water molecules are polar, having regions of partial

relied heavily on biochemistry and molecular biology, the chemistry and

positive and partial negative charge that permit them to form hydrogen bonds with one another and with other charged substances.

physics of the molecules that constitute living things. A biochemist may

2.6 Acids are hydrogen ion donors; bases are hydrogen ion acceptors. The pH scale is a convenient measure of the hydrogen ion concentration of a solution.

investigate the precise interactions among a cell’s atoms and molecules that maintain the energy flow essential to life, and a molecular biologist may study how proteins interact with deoxyribonucleic acid (DNA) in ways that control the expression of certain genes. However, an understanding of chemistry is essential to all biologists. An evolutionary biologist may

study evolutionary relationships by comparing the DNA of different types of organisms. An ecologist may study how energy is transferred among the organisms living in an estuary or monitor the biological effects of changes in the salinity of the water. A botanist may study unique compounds produced by plants and may even be a “chemical prospector,” seeking new sources of medicinal agents. In this chapter we lay a foundation for understanding how the structure of atoms determines the way they form chemical bonds to produce complex compounds. Most of our discussion focuses on small, simple substances known as inorganic compounds. Among the biologically important groups of inorganic compounds are water, many simple acids and bases, and simple salts. We pay particular attention to water, the most abundant substance in organisms and on Earth’s surface, and we examine how its unique properties affect living things as well as their nonliving environment. In Chapter 3 we extend our discussion to organic compounds, carbon-containing

particles. Physicists have discovered a number of subatomic particles, but for our purposes we need consider only three: electrons, protons, and neutrons. An electron is a particle that carries a unit of negative electric charge; a proton carries a unit of positive charge; and a neutron is an uncharged particle. In an electrically neutral atom, the number of electrons is equal to the number of protons. Clustered together, protons and neutrons compose the atomic nucleus. Electrons, however, have no fixed locations and move rapidly through the mostly empty space surrounding the atomic nucleus.

An atom is uniquely identified by its number of protons Every element has a fixed number of protons in the atomic nucleus, known as the atomic number. It is written as a subscript to the left of the chemical symbol. Thus, 1H indicates that the hydrogen

compounds that are generally large and complex. In all but the simplest organic compounds, two or more carbon atoms are bonded to

TABLE 2-1

Functions of Elements in Organisms

each other to form the backbone, or skeleton, of the molecule. Element* (chemical symbol)

2.1

ELEMENTS AND ATOMS

■ ■ LEARNING OBJECTIVES 1 2

3

Name the principal chemical elements in living things and provide an important function of each. Compare the physical properties (mass and charge) and locations of electrons, protons, and neutrons. Distinguish between the atomic number and the mass number of an atom. Define the terms orbital and electron shell. Relate electron shells to principal energy levels.

Elements are substances that cannot be broken down into simpler substances by ordinary chemical reactions. Each element has a chemical symbol: usually the first letter or first and second letters of the English or Latin name of the element. For example, O is the symbol for oxygen, C for carbon, H for hydrogen, N for nitrogen, and Na for sodium (from the Latin word natrium). Just four elements—oxygen, carbon, hydrogen, and nitrogen—are responsible for more than 96% of the mass of most organisms. Others, such as calcium, phosphorus, potassium, and magnesium, are also consistently present but in smaller quantities. Some elements, such as iodine and copper, are known as trace elements because they are required only in minute amounts. TABLE 2-1 lists the elements that make up organisms and briefly explains the importance of each in typical plants and animals. An atom is defined as the smallest portion of an element that retains its chemical properties. Atoms are much too small to be visible under a light microscope. However, by sophisticated techniques (such as scanning tunneling microscopy, with magnifications as great as 5 million times) researchers have been able to photograph the positions of some large atoms in molecules. The components of atoms are tiny particles of matter (anything that has mass and takes up space) known as subatomic

O

Oxygen

C

Carbon

H

Hydrogen

N

Nitrogen

Ca

Calcium

P

Phosphorus

K

Potassium

S Na

Sulfur Sodium

Mg

Magnesium

Cl

Chlorine

Fe

Iron

Functions Required for cellular respiration; present in most organic compounds; component of water Forms backbone of organic molecules; each carbon atom can form four bonds with other atoms Present in most organic compounds; component of water; hydrogen ion (H+) is involved in some energy transfers Component of proteins and nucleic acids; component of chlorophyll in plants Structural component of bones and teeth; calcium ion (Ca2+) is important in muscle contraction, conduction of nerve impulses, and blood clotting; associated with plant cell wall Component of nucleic acids and of phospholipids in membranes; important in energy transfer reactions; structural component of bone Potassium ion (K+) is a principal positive ion (cation) in interstitial (tissue) fluid of animals; important in nerve function; affects muscle contraction; controls opening of stomata in plants Component of most proteins Sodium ion (Na+) is a principal positive ion (cation) in interstitial (tissue) fluid of animals; important in fluid balance; essential for conduction of nerve impulses; important in photosynthesis in plants Needed in blood and other tissues of animals; activates many enzymes; component of chlorophyll in plants Chloride ion (Cl−) is principal negative ion (anion) in interstitial (tissue) fluid of animals; important in water balance; essential for photosynthesis Component of hemoglobin in animals; activates certain enzymes

*Other elements found in very small (trace) amounts in animals, plants, or both include iodine (I), manganese (Mn), copper (Cu), zinc (Zn), cobalt (Co), fluorine (F), molybdenum (Mo), selenium (Se), boron (B), silicon (Si), and a few others.

nucleus contains 1 proton, and 8O means that the oxygen nucleus contains 8 protons. The atomic number determines an atom’s identity and defines the element. The periodic table is a chart of the elements arranged in order by atomic number (FIG. 2-1 and Appendix A). The periodic table is useful because it lets us simultaneously correlate many of the relationships among the various elements. Figure 2-1 includes representations of the electron configurations of several elements important in organisms. These Bohr models, which show the electrons arranged in a series of concentric circles around the nucleus, are convenient to use, but inaccurate. The space outside the nucleus is actually extremely large compared

KEY POINT

to the nucleus, and as you will see, electrons do not actually circle the nucleus in fixed concentric pathways.

Protons plus neutrons determine atomic mass The mass of a subatomic particle is exceedingly small, much too small to be conveniently expressed in grams or even micrograms.1 Such masses are expressed in terms of the atomic mass unit (amu), also called the dalton in honor of John Dalton, the English 1

Tables of commonly used units of scientific measurement are printed inside the back cover of this text.

The periodic table provides information about the elements: their compositions, structures, and chemical behavior.

Chemical symbol

O

Atomic number H

1

HYDROGEN

8

OXYGEN

Chemical name

N

7

NITROGEN

Number of e– in each energy level

Mg

12

C

MAGNESIUM

2•6

6

AT. MASS 16.00 amu

CARBON

1 AT. MASS 1.01 amu

2•5

H

Na

He

AT. MASS 14.01 amu

Ne

11

10

NEON

SODIUM

Li

Be

B

C

N

O

F

Ne

Al

Si

P

S

Cl

Ar

Se

Br

Kr

2•8•2

Na Mg K

2•4

AT. MASS 24.31 amu

AT. MASS 12.01 amu

Ca Sc

Ti

V

Cr

Mn Fe

Co Ni

Cu Zn

Ga Ge As

2•8 AT. MASS 20.18 amu

2•8•1 AT. MASS 22.99 amu

K

19

POTASSIUM

Rb Sr

Y

Zr

Nb Mo Tc

Cs

(L)

Hf

Ta

Fr

2•8•8•1 AT. MASS 39.10 amu

Ba

Ca

Ra (A)

(L) (A)

La

W

Ru Rh Pd

Re Os Ir

Pt

Ag

Sn

Cd In

P

15

PHOSPHORUS

Sb

Te

I

Xe

Bi

Po

At

Rn

Cl S

20

16

SULFUR

CALCIUM

C

Sm Eu

Gd

2•8•7

Tm Y

2•8•5

AT. MASS 35.45 amu

AT. MASS 30.97 amu

Ac

17

CHLORINE

T

2•8•8•2

Pu

Am Cm Bk

Cf

Es

Fm Md N

2•8•6 AT. MASS 32.07 amu

AT. MASS 40.08 amu

FIGURE 2-1 The periodic table Note the Bohr models depicting the electron configuration of atoms of some biologically important elements, plus neon, which is unreactive, because its valence shell is full (to be discussed later in the chapter).

Although the Bohr model does not depict electron configurations accurately, it is commonly used because of its simplicity and convenience. A complete periodic table is given in Appendix A.

Particle

Charge

Approximate Mass

Location

Proton Neutron Electron

Positive Neutral Negative

1 amu 1 amu Approx. 1/1800 amu

Nucleus Nucleus Outside nucleus

Isotopes of an element differ in number of neutrons Most elements consist of a mixture of atoms with different numbers of neutrons and thus different masses. Such atoms are called isotopes. Isotopes of the same element have the same number of protons and electrons; only the number of neutrons varies. The three isotopes of hydrogen, 11H (ordinary hydrogen), 21H (deuterium), and 31H (tritium), contain 0, 1, and 2 neutrons, respectively. FIGURE 2-2 shows Bohr models of two isotopes of carbon, 12 14 6 C and 6 C. The mass of an element is expressed as an average of the masses of its isotopes (weighted by their relative abundance in nature). For example, the atomic mass of hydrogen is not 1.0 amu, but 1.0079 amu, reflecting the natural occurrence of small amounts of deuterium and tritium in addition to the more abundant ordinary hydrogen. Because they have the same number of electrons, all isotopes of a given element have essentially the same chemical characteristics. However, some isotopes are unstable and tend to break down, or decay, to a more stable isotope (usually becoming a different element); such radioisotopes emit radiation when they decay. For example, the radioactive decay of 146 C occurs as a neutron decomposes to form a proton and a fast-moving electron, which is emitted from the atom as a form of radiation known as a beta (b) particle. The resulting stable atom is the common form of nitrogen, 147 N. Using sophisticated instruments, scientists can detect and measure b particles and other types of radiation. Radioactive decay can also be detected by a method known as autoradiography, in which radiation causes the appearance of dark silver grains in photographic film (FIG. 2-3). 2

Unlike weight, mass is independent of the force of gravity. For convenience, however, we consider mass and weight equivalent. Atomic weight has the same numerical value as atomic mass, but it has no units.

–

–

+ –

–

+

+

+ +

–

–

+ +

–

–

+

+ +

–

–

+

+

–

–

Carbon-14 ( 146C) (6p, 8n)

Carbon-12 ( 126C) (6p, 6n)

FIGURE 2-2 Isotopes Carbon-12 (126 C) is the most common isotope of carbon. Its nucleus contains 6 protons and 6 neutrons, so its atomic mass is 12. Carbon-14 (146 C) is a rare radioactive carbon isotope. It contains 8 neutrons, so its atomic mass is 14.

Because the different isotopes of a given element have the same chemical characteristics, they are essentially interchangeable in molecules. Molecules containing radioisotopes are usually metabolized and/or localized in the organism in a similar way to their nonradioactive counterparts, and they can be substituted. For this reason, radioisotopes such as 3H (tritium), 14C, and 32P are extremely valuable research tools used, for example, in dating fossils (see Fig. 18-10), tracing biochemical pathways, determining the sequence of genetic information in DNA (see Fig. 15-10), and understanding sugar transport in plants. In medicine, radioisotopes are used for both diagnosis and treatment. The location and/or metabolism of a substance such as a hormone or drug can be followed in the body by labeling the substance with a radioisotope such as carbon-14 or tritium. Radioisotopes are used to test thyroid gland function, to provide images of blood flow in the arteries supplying the cardiac muscle, and to

Peter J. Bryant/Biological Photo Service

chemist who formulated an atomic theory in the early 1800s. One amu is equal to the approximate mass of a single proton or a single neutron. Protons and neutrons make up almost all the mass of an atom. The mass of a single electron is only about 1/1800 the mass of a proton or neutron. The atomic mass of an atom is a number that indicates approximately how much matter it contains compared with another atom. This value is determined by adding the number of protons to the number of neutrons and expressing the result in atomic mass units or daltons.2 The mass of the electrons is ignored because it is so small. The atomic mass number is indicated by a superscript to the left of the chemical symbol. The common form of the oxygen atom, with 8 protons and 8 neutrons in its nucleus, has an atomic number of 8 and a mass of 16 amu. It is indicated by the symbol 168 O . The characteristics of protons, electrons, and neutrons are summarized in the following table:

Concentrated silver grains

50 μm

FIGURE 2-3 Autoradiography The chromosomes of the fruit fly, Drosophila melanogaster, shown in this light micrograph, have been covered with photographic film in which silver grains (dark spots) are produced when tritium (3H) that has been incorporated into DNA undergoes radioactive decay. The concentrations of silver grains (arrows) mark the locations of specific DNA molecules.

study many other aspects of body function and chemistry. Because radiation can interfere with cell division, radioisotopes have been used therapeutically in treating cancer, a disease often characterized by rapidly dividing cells.

Electrons move in orbitals corresponding to energy levels Electrons move through characteristic regions of 3-D space, or orbitals. Each orbital contains a maximum of 2 electrons. Because

KEY POINT

it is impossible to know an electron’s position at any given time, orbitals are most accurately depicted as “electron clouds,” shaded areas whose density is proportional to the probability that an electron is present there at any given instant. The energy of an electron depends on the orbital it occupies. Electrons in orbitals with similar energies, said to be at the same principal energy level, make up an electron shell (FIG. 2-4). In general, electrons in a shell with a greater average distance from the nucleus have greater energy than those in a shell close to the nucleus. The reason is that energy is required to move a nega-

Electrons occupy orbitals corresponding to energy levels.

z

Nucleus

y 1s

(a) The first principal energy level contains a maximum of 2 electrons, occupying a single spherical orbital (designated 1s). The electrons depicted in the diagram could be present anywhere in the blue area.

x

z

z

z

2px

2s

z

2py

2pz

y

y

y

y

x

x

x

x

(b) The second principal energy level includes four orbitals, each with a maximum of 2 electrons: one spherical (2s) and three dumbbell-shaped (2p) orbitals at right angles to one another. z

1s

2s y 2py 2px x

2pz

(c) Orbitals of the first and second principal energy levels of a neon atom are shown superimposed. Note that the single 2s orbital plus three 2p orbitals make up neon's full valence shell of 8 electrons. Compare this more realistic view of the atomic orbitals with the Bohr model of a neon atom at right.

(d) Neon atom (Bohr model)

FIGURE 2-4 Animated Atomic orbitals Each orbital is represented as an “electron cloud.” The arrows labeled x, y, and z establish the imaginary axes of the atom.

tively charged electron farther away from the positively charged nucleus. The most energetic electrons, known as valence electrons, are said to occupy the valence shell. The valence shell is represented as the outermost concentric ring in a Bohr model. As we will see in the next sections, it is these valence electrons that play a key role in chemical reactions. An electron can move to an orbital farther from the nucleus by receiving more energy, or it can give up energy and sink to a lower energy level in an orbital nearer the nucleus. Changes in electron energy levels are important in energy conversions in organisms. For example, during photosynthesis, light energy absorbed by chlorophyll molecules causes electrons to move to a higher energy level (see Fig. 9-3).

Review ■

■

■

Do all atoms of an element have the same atomic number? the same atomic mass? What is a radioisotope? What are some ways radioisotopes are used in biological research? How do electrons in different orbitals of the same electron shell compare with respect to their energy?

2.2 CHEMICAL REACTIONS ■ ■ LEARNING OBJECTIVES 4 Explain how the number of valence electrons of an atom is related 5 6

to its chemical properties. Distinguish among simplest, molecular, and structural chemical formulas. Explain why the mole concept is so useful to chemists.

The chemical behavior of an atom is determined primarily by the number and arrangement of its valence electrons. The valence shell of hydrogen or helium is full (stable) when it contains 2 electrons. The valence shell of any other atom is full when it contains 8 electrons. When the valence shell is not full, the atom tends to lose, gain, or share electrons to achieve a full outer shell. The valence shells of all isotopes of an element are identical; for this reason, they have similar chemical properties and can substitute for one another in chemical reactions (for example, tritium can substitute for ordinary hydrogen). Elements in the same vertical column (belonging to the same group) of the periodic table have similar chemical properties because their valence shells have similar tendencies to lose, gain, or share electrons. For example, chlorine and bromine, included in a group commonly known as the halogens, are highly reactive. Because their valence shells have 7 electrons, they tend to gain an electron in chemical reactions. By contrast, hydrogen, sodium, and potassium each have a single valence electron, which they tend to give up or share with another atom. Helium (He) and neon (Ne) belong to a group referred to as the noble gases. They are quite unreactive because their valence shells are full. Notice in Figure 2-1 the incomplete valence shells of some of the elements important in organisms, including carbon, hydrogen, oxygen, and nitrogen, and compare them with the full valence shell of neon in Figure 2-4d.

Atoms form compounds and molecules Two or more atoms may combine chemically. When atoms of different elements combine, the result is a chemical compound. A chemical compound consists of atoms of two or more different elements combined in a fixed ratio. For example, water is a chemical compound composed of hydrogen and oxygen in a ratio of 2:1. Common table salt, sodium chloride, is a chemical compound made up of sodium and chlorine in a 1:1 ratio. Two or more atoms may become joined very strongly to form a stable particle called a molecule. For example, when two atoms of oxygen combine chemically, a molecule of oxygen is formed. Water is a molecular compound, with each molecule consisting of two atoms of hydrogen and one of oxygen. However, as you will see, not all compounds are made up of molecules. Sodium chloride (common table salt) is an example of a compound that is not molecular.

Simplest, molecular, and structural chemical formulas give different information A chemical formula is a shorthand expression that describes the chemical composition of a substance. Chemical symbols indicate the types of atoms present, and subscript numbers indicate the ratios among the atoms. There are several types of chemical formulas, each providing specific kinds of information. In a simplest formula (also known as an empirical formula), the subscripts give the smallest whole-number ratios for the atoms present in a compound. For example, the simplest formula for hydrazine is NH2, indicating a 1:2 ratio of nitrogen to hydrogen. (Note that when a single atom of a type is present, the subscript number 1 is never written.) In a molecular formula, the subscripts indicate the actual numbers of each type of atom per molecule. The molecular formula for hydrazine is N2H4, which indicates that each molecule of hydrazine consists of two atoms of nitrogen and four atoms of hydrogen. The molecular formula for water, H2O, indicates that each molecule consists of two atoms of hydrogen and one atom of oxygen. A structural formula shows not only the types and numbers of atoms in a molecule but also their arrangement. For example, the structural formula for water is HOOOH. As you will learn in Chapter 3, it is common for complex organic molecules with different structural formulas to share the same molecular formula.

One mole of any substance contains the same number of units The molecular mass of a compound is the sum of the atomic masses of the component atoms of a single molecule; thus, the molecular mass of water, H2O, is (hydrogen: 2 × 1 amu) + (oxygen: 1 × 16 amu), or 18 amu. (Because of the presence of isotopes, atomic mass values are not whole numbers, but for easy calculation each atomic mass value has been rounded off to a whole number.) Similarly, the molecular mass of glucose (C6H12O6), a simple sugar that is a key compound in cell metabolism, is (carbon: 6 × 12 amu) + (hydrogen: 12 × 1 amu) + (oxygen: 6 × 16 amu), or 180 amu.

The amount of an element or compound whose mass in grams is equivalent to its atomic or molecular mass is 1 mole (mol). Thus, 1 mol of water is 18 grams (g), and 1 mol of glucose has a mass of 180 g. The mole is an extremely useful concept because it lets us make meaningful comparisons between atoms and molecules of very different mass. The reason is that 1 mol of any substance always has exactly the same number of units, whether those units are small atoms or large molecules. The very large number of units in a mole, 6.02 × 1023, is known as Avogadro’s number, named for the Italian physicist Amadeo Avogadro, who first calculated it. Thus, 1 mol (180 g) of glucose contains 6.02 × 1023 molecules, as does 1 mol (2 g) of molecular hydrogen (H2). Although it is impossible to count atoms and molecules individually, a scientist can calculate them simply by weighing a sample. Molecular biologists usually deal with smaller values, either millimoles (mmol, one-thousandth of a mole) or micromoles (mmol, one-millionth of a mole). The mole concept also lets us make useful comparisons among solutions. A 1 molar solution, represented by 1 M, contains 1 mol of that substance dissolved in a total volume of 1 liter (L). For example, we can compare 1 L of a 1 M solution of glucose with 1 L of a 1 M solution of sucrose (table sugar, a larger molecule). They differ in the mass of the dissolved sugar (180 g and 340 g, respectively), but they each contain 6.02 × 1023 sugar molecules.

Review ■

■ ■

Why is a radioisotope able to substitute for an ordinary (nonradioactive) atom of the same element in a molecule? Which kind of chemical formula provides the most information? How many particles would be included in 1 g of hydrogen atoms? in 2 g of hydrogen molecules?

2.3 CHEMICAL BONDS ■ ■ LEARNING OBJECTIVE 7

Distinguish among covalent bonds, ionic bonds, hydrogen bonds, and van der Waals interactions. Compare them in terms of the mechanisms by which they form and their relative strengths.

Atoms can be held together by forces of attraction called chemical bonds. Each bond represents a certain amount of chemical energy. Bond energy is the energy necessary to break a chemical bond. The valence electrons dictate how many bonds an atom can form. The two principal types of strong chemical bonds are covalent bonds and ionic bonds.

In covalent bonds electrons are shared Chemical equations describe chemical reactions During any moment in the life of an organism—a bacterial cell, a mushroom, or a butterfly—many complex chemical reactions are taking place. Chemical reactions, such as the reaction between glucose and oxygen, can be described by means of chemical equations: C6 H12 O6 + 6 O2 ¡ Glucose

Oxygen

6 CO2 Carbon dioxide

+ 6 H2O + energy Water

In a chemical equation, the reactants, the substances that participate in the reaction, are generally written on the left side, and the products, the substances formed by the reaction, are written on the right side. The arrow means “yields” and indicates the direction in which the reaction proceeds. Chemical compounds react with one another in quantitatively precise ways. The numbers preceding the chemical symbols or formulas (known as coefficients) indicate the relative number of atoms or molecules reacting. For example, 1 mol of glucose burned in a fire or metabolized in a cell reacts with 6 mol of oxygen to form 6 mol of carbon dioxide and 6 mol of water. Many reactions can proceed simultaneously in the reverse direction (to the left) and the forward direction (to the right). At dynamic equilibrium, the rates of the forward and reverse reactions are equal (see Chapter 7). Reversible reactions are indicated by double arrows: CO2 Carbon dioxide

+

H2O

H2CO3

Water

Carbonic acid

In this example, the arrows are drawn in different lengths to indicate that when the reaction reaches equilibrium, there will be more reactants (CO2 and H2O) than product (H2CO3).

Covalent bonds involve the sharing of electrons between atoms in a way that results in each atom having a filled valence shell. A molecule consists of atoms joined by covalent bonds. A simple example of a covalent bond is the joining of two hydrogen atoms in a molecule of hydrogen gas, H2. Each atom of hydrogen has 1 electron, but 2 electrons are required to complete its valence shell. The hydrogen atoms have equal capacities to attract electrons, so neither donates an electron to the other. Instead, the two hydrogen atoms share their single electrons so that the 2 electrons are attracted simultaneously to the 2 protons in the two hydrogen nuclei. The 2 electrons whirl around both atomic nuclei, thus forming the covalent bond that joins the two atoms. Similarly, unlike atoms can also be linked by covalent bonds to form molecules; the resulting compound is a covalent compound. A simple way of representing the electrons in the valence shell of an atom is to use dots placed around the chemical symbol of the element. Such a representation is called the Lewis structure of the atom, named for G. N. Lewis, an American chemist who developed this type of notation. In a water molecule, two hydrogen atoms are covalently bonded to an oxygen atom: H +H + O

H O H

Oxygen has 6 valence electrons; by sharing electrons with two hydrogen atoms, it completes its valence shell of 8. At the same time, each hydrogen atom obtains a complete valence shell of 2. (Note that in the structural formula HOOOH, each pair of shared electrons constitutes a covalent bond, represented by a solid line. Unshared electrons are usually omitted in a structural formula.) The carbon atom has 4 electrons in its valence shell, all of which are available for covalent bonding:

C

When one carbon and four hydrogen atoms share electrons, a molecule of the covalent compound methane, CH4, is formed: H H C H H

H or

H

C

H

H

Lewis structure

Structural formula

The nitrogen atom has 5 electrons in its valence shell. Recall that each orbital can hold a maximum of 2 electrons. Usually, 2 electrons occupy one orbital, leaving 3 electrons available for sharing with other atoms: N

When a nitrogen atom shares electrons with three hydrogen atoms, a molecule of the covalent compound ammonia, NH3, is formed: H N H H

or

H

N

H

H

Lewis structure

Structural formula

When one pair of electrons is shared between two atoms, the covalent bond is called a single covalent bond (FIG. 2-5a). Two hydrogen atoms share a single pair of electrons. Two oxygen atoms

KEY POINT

may achieve stability by forming covalent bonds with each other. Each oxygen atom has 6 electrons in its outer shell. To become stable, the two atoms share two pairs of electrons, forming molecular oxygen (FIG. 2-5b). When two pairs of electrons are shared in this way, the covalent bond is called a double covalent bond, which is represented by two parallel solid lines. Similarly, a triple covalent bond is formed when three pairs of electrons are shared between two atoms (represented by three parallel solid lines). The number of covalent bonds usually formed by the atoms in biologically important molecules is summarized as follows: Atom Hydrogen Oxygen Carbon Nitrogen Phosphorus Sulfur

Symbol

Covalent Bonds

H O C N P S

1 2 4 3 5 2

The function of a molecule is related to its shape In addition to being composed of atoms with certain properties, each kind of molecule has a characteristic size and a general overall shape. Although the shape of a molecule may change (within certain limits), the functions of molecules in living cells are dictated largely by their geometric shapes. A molecule that consists of two atoms is linear. Molecules composed of more than two atoms may

Covalent bonds form when atoms share electrons.

+

Hydrogen (H)

H H

Hydrogen (H)

Molecular hydrogen (H2)

or

H H

(a) Single covalent bond formation. Two hydrogen atoms achieve stability by sharing a pair of electrons, thereby forming a molecule of hydrogen. In the structural formula on the right, the straight line between the hydrogen atoms represents a single covalent bond.

+

Oxygen (O)

Oxygen (O)

Molecular oxygen (O2) (double bond is formed)

or

O

O

O

O

(b) Double covalent bond formation. In molecular oxygen, two oxygen atoms share two pairs of electrons, forming a double covalent bond. The parallel straight lines in the structural formula represent a double covalent bond.

FIGURE 2-5 Electron sharing in covalent compounds

H

H

C

H H

Methane (CH4)

FIGURE 2-6 Orbital hybridization in methane The four hydrogens are located at the corners of a tetrahedron because of hybridization of the valence shell orbitals of carbon.

have more complicated shapes. The geometric shape of a molecule provides the optimal distance between the atoms to counteract the repulsion of electron pairs. When an atom forms covalent bonds with other atoms, the orbitals in the valence shell may become rearranged in a process known as orbital hybridization, thereby affecting the shape of the resulting molecule. For example, when four hydrogen atoms combine with a carbon atom to form a molecule of methane (CH4), the hybridized valence shell orbitals of the carbon form a geometric structure known as a tetrahedron, with one hydrogen atom present at each of its four corners (FIG. 2-6; see also Fig. 3-2a). We will explore the importance of molecular shape in more detail in Chapter 3 and in our discussion of the properties of water in this chapter. Covalent bonds can be nonpolar or polar Atoms of different elements vary in their affinity for electrons. Electronegativity is a measure of an atom’s attraction for shared electrons in chemical bonds. Very electronegative atoms such as oxygen, nitrogen, fluorine, and chlorine are sometimes called “electron greedy.” When covalently bonded atoms have similar electronegativities, the electrons are shared equally and the covalent bond is described as nonpolar. The covalent bond of the hydrogen molecule is nonpolar, as are the covalent bonds of molecular oxygen and methane.

In a covalent bond between two different elements, such as oxygen and hydrogen, the electronegativities of the atoms may be different. If so, electrons are pulled closer to the atomic nucleus of the element with the greater electron affinity (in this case, oxygen). A covalent bond between atoms that differ in electronegativity is called a polar covalent bond. Such a bond has two dissimilar ends (or poles), one with a partial positive charge and the other with a partial negative charge. Each of the two covalent bonds in water is polar because there is a partial positive charge at the hydrogen end of the bond and a partial negative charge at the oxygen end, where the “shared” electrons are more likely to be. Covalent bonds differ in their degree of polarity, ranging from those in which the electrons are equally shared (as in the nonpolar hydrogen molecule) to those in which the electrons are much closer to one atom than to the other (as in water). Oxygen is quite electronegative and forms polar covalent bonds with carbon, hydrogen, and many other atoms. Nitrogen is also strongly electronegative, although less so than oxygen. A molecule with one or more polar covalent bonds can be polar even though it is electrically neutral as a whole. The reason is that a polar molecule has one end with a partial positive charge and another end with a partial negative charge. One example is water (FIG. 2-7). The polar bonds between the hydrogens and the oxygen are arranged in a V shape, rather than linearly. The oxygen end constitutes the negative pole of the molecule, and the end with the two hydrogens is the positive pole.

Ionic bonds form between cations and anions Some atoms or groups of atoms are not electrically neutral. A particle with 1 or more units of electric charge is called an ion. An atom becomes an ion if it gains or loses 1 or more electrons. An atom with 1, 2, or 3 electrons in its valence shell tends to lose electrons to other atoms. Such an atom then becomes positively charged because its nucleus contains more protons than the number of electrons orbiting around the nucleus. These positively charged ions are termed cations. Atoms with 5, 6, or 7 valence electrons tend to gain electrons from other atoms and become negatively charged anions. The properties of ions are quite different from those of the electrically neutral atoms from which they were derived. For example, although chlorine gas is a poison, chloride ions (Cl−) are Oxygen part Partial negative charge at oxygen end of molecule

Hydrogen (H)

Oxygen (O)

+

– Hydrogen parts

–

Hydrogen (H)

+ Water molecule (H2O)

FIGURE 2-7 Water, a polar molecule Note that the electrons tend to stay closer to the nucleus of the oxygen atom than to the hydrogen nuclei. This results in a partial negative charge on the oxygen portion of the molecule and a partial positive charge at the hydrogen end. Although the water molecule as a whole is electrically neutral, it is a polar covalent compound.

Partial positive charge at hydrogen end of molecule

17 protons

11 protons

Muscle fiber and

Bloom and Fawcett Textbook of Technology

Nerve

11 electrons Sodium (Na)

17 electrons Chlorine (Cl)

+

–

100 μm

FIGURE 2-8 Ions and biological processes Sodium, potassium, and chloride ions are essential for this nerve cell to stimulate these muscle fibers, initiating a muscle contraction. Calcium ions in the muscle cell are required for muscle contraction.

essential to life (see Table 2-1). Because their electric charges provide a basis for many interactions, cations and anions are involved in energy transformations within the cell, the transmission of nerve impulses, muscle contraction, and many other biological processes (FIG. 2-8). A group of covalently bonded atoms can also become an ion (polyatomic ion). Unlike a single atom, a group of atoms can lose or gain protons (derived from hydrogen atoms) as well as electrons. Therefore, a group of atoms can become a cation if it loses 1 or more electrons or gains 1 or more protons. A group of atoms becomes an anion if it gains 1 or more electrons or loses 1 or more protons. An ionic bond forms as a consequence of the attraction between the positive charge of a cation and the negative charge of an anion. An ionic compound is a substance consisting of anions and cations bonded by their opposite charges. A good example of how ionic bonds are formed is the attraction between sodium ions and chloride ions. A sodium atom has 1 electron in its valence shell. It cannot fill its valence shell by obtaining 7 electrons from other atoms because it would then have a large unbalanced negative charge. Instead, it gives up its single valence electron to a very electronegative atom, such as chlorine, which acts as an electron acceptor (FIG. 2-9). Chlorine cannot give up the 7 electrons in its valence shell because it would then have a large positive charge. Instead, it strips an electron from an electron donor (sodium, in this example) to complete its valence shell. When sodium reacts with chlorine, sodium’s valence electron is transferred completely to chlorine. Sodium becomes a cation, with 1 unit of positive charge (Na+). Chlorine becomes an anion, a chloride ion with 1 unit of negative charge (Cl−). These ions attract each other as a result of their opposite charges. This electrical attraction in ionic bonds holds them together to form NaCl, sodium chloride, or common table salt.

10 electrons Sodium ion (Na+)

18 electrons Chloride ion (Cl–)

Sodium chloride (NaCl)

Cl– Na+ Cl– Na+

Cl– Na+ Na+

Arrangement of atoms in a crystal of salt

FIGURE 2-9 Animated Ionic bonding Sodium becomes a positively charged ion when it donates its single valence electron to chlorine, which has 7 valence electrons. With this additional electron, chlorine completes its valence shell and becomes a negatively charged chloride ion. These sodium and chloride ions are attracted to one another by their unlike electric charges, forming the ionic compound sodium chloride.

The term molecule does not adequately explain the properties of ionic compounds such as NaCl. When NaCl is in its solid crystal state, each ion is actually surrounded by six ions of opposite charge. The simplest formula, NaCl, indicates that sodium ions and chloride ions are present in a 1:1 ratio, but the actual crystal has no discrete molecules composed of one Na+ and one Cl− ion. Compounds joined by ionic bonds, such as sodium chloride, have a tendency to dissociate (separate) into their individual ions when placed in water: NaCl Sodium chloride

in H2O

¡

Na+ Sodium ion

+

Cl– Chloride ion

In the solid form of an ionic compound (that is, in the absence of water), ionic bonds are very strong. Water, however, is an excellent solvent; as a liquid it is capable of dissolving many substances, particularly those that are polar or ionic, because of the polarity of water molecules. The localized partial positive charge (on the hydrogen atoms) and partial negative charge (on the oxygen atom) on each water molecule attract and surround the anions and cations, respectively, on the surface of an ionic solid. As a result, the solid dissolves. A dissolved substance is referred to as a solute. In solution, each cation and anion of the ionic compound is surrounded by oppositely charged ends of the water molecules. This process is known as hydration (FIG. 2-10). Hydrated ions still interact with one another to some extent, but the transient ionic bonds formed are much weaker than those in a solid crystal.

Hydrogen bonds are weak attractions Another type of bond important in organisms is the hydrogen bond. When hydrogen combines with oxygen (or with another relatively electronegative atom such as nitrogen), it acquires a partial positive charge because its electron spends more time closer to the electronegative atom. Hydrogen bonds tend to form between an atom with a partial negative charge and a hydrogen atom that is covalently bonded to oxygen or nitrogen (FIG. 2-11). The atoms involved may be in two parts of the same large molecule or in two different molecules. Water molecules interact with one another extensively through hydrogen bond formation.

Salt

Electronegative atoms O

H

+

–

H N

H

H Hydrogen bond

H

FIGURE 2-11 Animated Hydrogen bonding A hydrogen bond (dotted line) can form between two molecules with regions of unlike partial charge. Here, the nitrogen atom of an ammonia molecule is joined by a hydrogen bond to a hydrogen atom of a water molecule.

Hydrogen bonds are readily formed and broken. Although individually relatively weak, hydrogen bonds are collectively strong when present in large numbers. Furthermore, they have a specific length and orientation. As you will see in Chapter 3, these features are very important in determining the 3-D structure of large molecules such as DNA and proteins.

van der Waals interactions are weak forces Even electrically neutral, nonpolar molecules can develop transient regions of weak positive and negative charge. These slight charges develop as a consequence of the fact that electrons are in constant motion. A region with a temporary excess of electrons will have a weak negative charge, whereas one with an electron deficit will have a weak positive charge. Adjacent molecules may interact in regions of slight opposite charge. These attractive forces, called van der Waals interactions, operate over very short distances and are weaker and less specific than the other types of interactions we have considered. They are most important when they occur in large numbers and when the shapes of the molecules permit close contact between the atoms. Although a single interaction is very weak, the binding force of a large number of these interactions working together can be significant.

Review –

–

–

O H H Cl– –

–

–

–

–

–

Na+ –

–

– – + Cl– + Cl– – Na Na+ Cl Cl–

–

–

H – H+ O –

■

+

–

Cl–

+ –

+

H– + OH

–

Na+

Cl–

–

–

H H+ O Na+

■ ■

O

+H

Are all compounds composed of molecules? Explain. What are the ways an atom or molecule can become an anion or a cation? How do ionic and covalent bonds differ? Under what circumstances can weak forces such as hydrogen bonds and van der Waals interactions play significant roles in biological systems?

–

O H H

–

–

Cl–

– –

–

–

+ H–

– –

■

–

–

–

2.4 REDOX REACTIONS ■ ■ LEARNING OBJECTIVE 8

FIGURE 2-10 Animated Hydration of an ionic compound When the crystal of NaCl is added to water, the sodium and chloride ions are pulled apart. When the NaCl is dissolved, each Na+ and Cl− is surrounded by water molecules electrically attracted to it.

Distinguish between the terms oxidation and reduction, and relate these processes to the transfer of energy.

Many energy conversions that go on in a cell involve reactions in which an electron transfers from one substance to another. The reason is that the transfer of an electron also involves the transfer

of the energy of that electron. Such an electron transfer is known as an oxidation–reduction, or redox reaction. Oxidation and reduction always occur together. Oxidation is a chemical process in which an atom, ion, or molecule loses one or more electrons. Reduction is a chemical process in which an atom, ion, or molecule gains one or more electrons. (The term refers to the fact that the gain of an electron results in the reduction of any positive charge that might be present.) Rusting—the combining of iron (symbol Fe) with oxygen— is a simple illustration of oxidation and reduction: 4 Fe + 3 O2 ¡

A large part of the mass of most organisms is water. In human tissues the percentage of water ranges from 20% in bones to 85% in brain cells; about 70% of our total body weight is water. As much as 95% of a jellyfish and certain plants is water. Water is the source, through photosynthesis, of the oxygen in the air we breathe, and its hydrogen atoms become incorporated into many organic compounds. Water is also the solvent for most biological reactions and a reactant or product in many chemical reactions. Water is important not only as an internal constituent of organisms but also as one of the principal environmental factors affecting them (FIG. 2-12). Many organisms live in the ocean or in freshwater rivers, lakes, or puddles. Water’s unique combination of physical and chemical properties is considered to have been essential to the origin of life as well as to the continued survival and evolution of life on Earth.

2 Fe 2O3 Iron (III) oxide

In rusting, each iron atom becomes oxidized as it loses 3 electrons. 4 Fe ¡ 4 Fe3+ + 12e− −

Hydrogen bonds form between water molecules

The e represents an electron; the + superscript in Fe represents an electron deficit. (When an atom loses an electron, it acquires 1 unit of positive charge from the excess of 1 proton. In our example, each iron atom loses 3 electrons and acquires 3 units of positive charge.) Recall that the oxygen atom is very electronegative, able to remove electrons from other atoms. In this reaction, oxygen becomes reduced when it accepts electrons from the iron. 3+

As discussed, water molecules are polar; that is, one end of each molecule bears a partial positive charge and the other a partial negative charge (see Fig. 2-7). The water molecules in liquid water and in ice associate by hydrogen bonds. The hydrogen atom of one water molecule, with its partial positive charge, is attracted to the oxygen atom of a neighboring water molecule, with its partial negative charge, forming a hydrogen bond. An oxygen atom in a water molecule has two regions of partial negative charge, and each of the two hydrogen atoms has a partial positive charge. Each water molecule can therefore form hydrogen bonds with a maximum of four neighboring water molecules (FIG. 2-13).

Review ■ ■

Why must oxidation and reduction occur simultaneously? Why are redox reactions important in some energy transfers?

2.5 WATER ■ ■ LEARNING OBJECTIVE

(a) Commonly known as "water bears," tardigrades, such as these members of the genus Echiniscus, are small animals (less than 1.2 mm long) that normally live in moist habitats, such as thin films of water on mosses.

100 μm

(b) When subjected to desiccation (dried out), tardigrades assume a barrel-shaped form known as a tun, remaining in this state, motionless but alive, for as long as 100 years. When rehydrated, they assume their normal appearance and activities.

Robert O. Schuster, courtesy of Diane R. Nelson

Redox reactions occur simultaneously because one substance must accept the electrons that are removed from the other. In a redox reaction, one component, the oxidizing agent, accepts 1 or more electrons and becomes reduced. Oxidizing agents other than oxygen are known, but oxygen is such a common one that its name was given to the process. Another reaction component, the reducing agent, gives up 1 or more electrons and becomes oxidized. In our example, there was a complete transfer of electrons from iron (the reducing agent) to oxygen (the oxidizing agent). Similarly, Figure 2-9 shows that an electron was transferred from sodium (the reducing agent) to chlorine (the oxidizing agent). Electrons are not easily removed from covalent compounds unless an entire atom is removed. In cells, oxidation often involves the removal of a hydrogen atom (an electron plus a proton that “goes along for the ride”) from a covalent compound; reduction often involves the addition of the equivalent of a hydrogen atom (see Chapter 7).

Diane R. Nelson

3 O2 + 12e− ¡ 6 O2−

10 μm

9 Explain how hydrogen bonds between adjacent water molecules govern many of the properties of water.

FIGURE 2-12 The effects of water on an organism

H O

H

– + H

H

+–

O

H H O

– +

H

+ –

H O Dennis Drenner

O

H

H

FIGURE 2-13 Hydrogen bonding of water molecules Each water molecule can form hydrogen bonds (dotted lines) with as many as four neighboring water molecules.

Water molecules have a strong tendency to stick to one another, a property known as cohesion. This is due to the hydrogen bonds among the molecules. Because of the cohesive nature of water molecules, any force exerted on part of a column of water is transmitted to the column as a whole. The major mechanism of water movement in plants (see Chapter 35) depends on the cohesive nature of water. Water molecules also display adhesion, the ability to stick to many other kinds of substances, most notably those with charged groups of atoms or molecules on their surfaces. These adhesive forces explain how water makes things wet. A combination of adhesive and cohesive forces accounts for capillary action, which is the tendency of water to move in narrow tubes, even against the force of gravity (FIG. 2-14). For example, water moves through the microscopic spaces between soil particles to the roots of plants by capillary action.

(a)

(b)

FIGURE 2-14 Capillary action (a) In a narrow tube, there is adhesion between the water molecules and the glass wall of the tube. Other water molecules inside the tube are then “pulled along” because of cohesion, which is due to hydrogen bonds between the water molecules. (b) In the wider tube, a smaller percentage of the water molecules line the glass wall. As a result, the adhesion is not strong enough to overcome the cohesion of the water molecules beneath the surface level of the container, and water in the tube rises only slightly.

FIGURE 2-15 Surface tension of water Hydrogen bonding between water molecules is responsible for the surface tension of water, which causes a dimpled appearance of the surface as these water striders (Gerris) walk across it. Fine hairs at the ends of the legs of these insects create highly water-repellent “cushions” of air.

Water has a high degree of surface tension because of the cohesion of its molecules, which have a much greater attraction for one another than for molecules in the air. Thus, water molecules at the surface crowd together, producing a strong layer as they are pulled downward by the attraction of other water molecules beneath them (FIG. 2-15).

Water molecules interact with hydrophilic substances by hydrogen bonding Because its molecules are polar, water is an excellent solvent, a liquid capable of dissolving many kinds of substances, especially polar and ionic compounds. Earlier we discussed how polar water molecules pull the ions of ionic compounds apart so that they dissociate (see Fig. 2-10). Because of its solvent properties and the tendency of the atoms in certain compounds to form ions in solution, water plays an important role in facilitating chemical reactions. Substances that interact readily with water are hydrophilic (“water-loving”). Examples include table sugar (sucrose, a polar compound) and table salt (NaCl, an ionic compound), which dissolve readily in water. Not all substances in organisms are hydrophilic, however. Many hydrophobic (“water-fearing”) substances found in living things are especially important because of their ability to form associations or structures that are not disrupted. Hydrophobic interactions occur between groups of nonpolar molecules. Such molecules are insoluble in water and tend to cluster together. This is not due to formation of bonds between the nonpolar molecules but rather to the fact that the hydrogen-bonded water molecules exclude them and in a sense “drive them together.” Hydrophobic interactions explain why oil tends to form globules when added to water. Examples of hydrophobic substances include fatty acids and cholesterol, discussed in Chapter 3.

Water helps maintain a stable temperature Hydrogen bonding explains the way water responds to changes in temperature. Water exists in three forms, which differ in their degree of hydrogen bonding: gas (vapor), liquid, and ice, a crystalline

Woodbridge Wilson/National Park Service

212°F

100°C

(a) Steam becoming water vapor (gas)

Gary R. Bonner

50°C

Barbara O’Donnell/Biological Photo Service

(b) Water (liquid)

32°F

0°C

(c) Ice (solid)

FIGURE 2-16 Animated Three forms of water (a) When water boils, as in this hot spring at Yellowstone National Park, many hydrogen bonds are broken, causing steam, consisting of minuscule water droplets, to form. If most of the remaining hydrogen bonds break, the molecules move more freely as water vapor (a gas). (b) Water molecules in a liquid state continually form, break, and re-form hydrogen bonds with one another. (c) In ice, each water molecule participates in four hydrogen bonds with adjacent molecules, resulting in a regular, evenly distanced crystalline lattice structure.

solid (FIG. 2-16). Hydrogen bonds are formed or broken as water changes from one state to another. Raising the temperature of a substance involves adding heat energy to make its molecules move faster, that is, to increase the energy of motion—kinetic energy—of the molecules (see Chapter 7). The term heat refers to the total amount of kinetic energy in a sample of a substance; temperature is a measure of the average kinetic energy of the particles. For the molecules to move more freely, some of the hydrogen bonds of water must be broken.

Much of the energy added to the system is used up in breaking the hydrogen bonds, and only a portion of the heat energy is available to speed the movement of the water molecules, thereby increasing the temperature of the water. Conversely, when liquid water changes to ice, additional hydrogen bonds must be formed, making the molecules less free to move and liberating a great deal of heat into the environment. Heat of vaporization, the amount of heat energy required to change 1 g of a substance from the liquid phase to the vapor phase,

is expressed in units called calories. A calorie (cal) is the amount of heat energy (equivalent to 4.184 joules [ J]) required to raise the temperature of 1 g of water 1 degree Celsius (C). Water has a high heat of vaporization—540 cal—because its molecules are held together by hydrogen bonds. The heat of vaporization of most other common liquid substances is much less. As a sample of water is heated, some molecules are moving much faster than others (they have more heat). These faster-moving molecules are more likely to escape the liquid phase and enter the vapor phase (see Fig. 2-16a). When they do, they take their heat with them, lowering the temperature of the sample, a process called evaporative cooling. For this reason, the human body can dissipate excess heat as sweat evaporates from the skin, and a leaf can keep cool in the bright sunlight as water evaporates from its surface. Hydrogen bonding is also responsible for water’s high specific heat; that is, the amount of energy required to raise the temperature of water is quite large. The specific heat of water is 1 cal/g of water per degree Celsius. Most other common substances, such as metals, glass, and ethyl alcohol, have much lower specific heat values. The specific heat of ethyl alcohol, for example, is 0.59 cal/ g/1°C (2.46 J/g/1°C). Because so much heat input is required to raise the temperature of water (and so much heat is lost when the temperature is lowered), the ocean and other large bodies of water have relatively constant temperatures. Thus, many organisms living in the ocean are provided with a relatively constant environmental temperature. The properties of water are crucial in stabilizing temperatures on Earth’s surface. Although surface water is only a thin film relative to Earth’s volume, the quantity is enormous compared to the exposed landmass. This relatively large mass of water resists both the warming effect of heat and the cooling effect of low temperatures. Hydrogen bonding causes ice to have unique properties with important environmental consequences. Liquid water expands as it freezes because the hydrogen bonds joining the water molecules in the crystalline lattice keep the molecules far enough apart to give ice a density about 10% less than the density of liquid water (see Fig. 2-16c). When ice has been heated enough to raise its temperature above 0°C (32°F), the hydrogen bonds are broken, freeing the molecules to slip closer together. The density of water is greatest at 4°C. Above that temperature water begins to expand again as the speed of its molecules increases. As a result, ice floats on the denser cold water. This unusual property of water has been important to the evolution of life. If ice had a greater density than water, the ice would sink; eventually, all ponds, lakes, and even the ocean would freeze solid from the bottom to the surface, making life impossible. When a deep body of water cools, it becomes covered with floating ice. The ice insulates the liquid water below it, retarding freezing and permitting organisms to survive below the icy surface. The high water content of organisms helps them maintain relatively constant internal temperatures. Such minimizing of temperature fluctuations is important because biological reactions can take place only within a relatively narrow temperature range.

Review ■

Why does water form hydrogen bonds?

■

■

What are some properties of water that result from hydrogen bonding? How do these properties contribute to the role of water as an essential component of organisms? How can weak forces, such as hydrogen bonds, have significant effects in organisms?

2.6 ACIDS, BASES, AND SALTS ■ ■ LEARNING OBJECTIVES 10 Contrast acids and bases, and discuss their properties. 11 Convert the hydrogen ion concentration (moles per liter) of a solution to a pH value and describe how buffers help minimize changes in pH.

12 Describe the composition of a salt and explain why salts are important in organisms.

Water molecules have a slight tendency to ionize, that is, to dissociate into hydrogen ions (H+) and hydroxide ions (OH−). The H+ immediately combines with a negatively charged region of a water molecule, forming a hydronium ion (H3O+). However, by convention, H+, rather than the more accurate H3O+, is used. In pure water, a small number of water molecules ionize. This slight tendency of water to dissociate is reversible as hydrogen ions and hydroxide ions reunite to form water. HOH

H+ + OH−

Because each water molecule splits into one hydrogen ion and one hydroxide ion, the concentrations of hydrogen ions and hydroxide ions in pure water are exactly equal (0.0000001 or 107 mol/L for each ion). Such a solution is said to be neutral, that is, neither acidic nor basic (alkaline). An acid is a substance that dissociates in solution to yield hydrogen ions (H+) and anions. Acid ¡ H+ + anion

An acid is a proton donor. (Recall that a hydrogen ion, or H+, is nothing more than a proton.) Hydrochloric acid (HCl) is a common inorganic acid. A base is defined as a proton acceptor. Most bases are substances that dissociate to yield a hydroxide ion (OH−) and a cation when dissolved in water. NaOH ¡ Na+ + OH− OH− + H+ ¡ H2O

A hydroxide ion can act as a base by accepting a proton (H+) to form water. Sodium hydroxide (NaOH) is a common inorganic base. Some bases do not dissociate to yield hydroxide ions directly. For example, ammonia (NH3) acts as a base by accepting a proton from water, producing an ammonium ion (NH4+) and releasing a hydroxide ion. NH3 + H2O ¡ NH4+ + OH−

pH is a convenient measure of acidity The degree of a solution’s acidity is generally expressed in terms of pH, defined as the negative logarithm (base 10) of the hydrogen ion concentration (expressed in moles per liter): pH = −log10[H+]

The brackets refer to concentration; therefore, the term [H+] means “the concentration of hydrogen ions,” which is expressed in moles per liter because we are interested in the number of hydrogen ions per liter. Because the range of possible pH values is broad, a logarithmic scale (with a 10-fold difference between successive units) is more convenient than a linear scale. Hydrogen ion concentrations are nearly always less than 1 mol/L. One gram of hydrogen ions dissolved in 1 L of water (a 1 M solution) may not sound impressive, but such a solution would be extremely acidic. The logarithm of a number less than 1 is a negative number; thus, the negative logarithm corresponds to a positive pH value. (Solutions with pH values less than zero can be produced but do not occur under biological conditions.) Whole-number pH values are easy to calculate. For instance, consider our example of pure water, which has a hydrogen ion concentration of 0.0000001 (10−7) mol/L. The logarithm is −7. The negative logarithm is 7; therefore, the pH is 7. TABLE 2-2 shows how to calculate pH values from hydrogen ion concentrations, and the reverse. For comparison, the table also includes the hydroxide ion concentrations, which can be calculated because the product of the hydrogen ion concentration and the hydroxide ion concentration is 1 × 10−14: [H+][OH−] = 1 × 10−14

Substance Gastric juice Pure water, neutral solution Household ammonia

Calculating pH Values and Hydroxide Ion Concentrations from Hydrogen Ion Concentrations [H+]* −2

pH

[OH−]†

Battery acid 0.0

1

Hydrochloric acid 0.8 Stomach acid 1.0

2

Stomach gastric juice 2.0

3

Vinegar 3.0

4 Beer 4.5 5

6

7

Black coffee 5.0

Rainwater 6.25 Cow milk 6.5 Distilled water 7.0 Blood 7.4

8

Seawater 8.0

9

Bleach 9.0

10

Increasing 11 alkalinity

Mono Lake, California 9.9

Household ammonia 11.5

13

Oven cleaner 13.0

14

Lye 14.0

−12

−2 −7

2 7

10 10−7

0.00000000001, 10−11

−11

11

10−3

[OH−] = hydroxide ion concentration (mol/L)

Increasing acidity

0

12

0.01, 10 0.0000001, 10−7

*[H+] = hydrogen ion concentration (mol/L) †

log [H+]

pH scale

Neutrality

Pure water is an example of a neutral solution; with a pH of 7, it has equal concentrations of hydrogen ions and hydroxide ions (the concentration of each is 10−7 mol/L). An acidic solution has a hydrogen ion concentration that is higher than its hydroxide ion concentration and has a pH value of less than 7. For example, the hydrogen ion concentration of a solution with pH 1 is 10 times that of a solution with pH 2. A basic solution has a hydrogen ion concentration that is lower than its hydroxide ion concentration and has a pH greater than 7. TABLE 2-2

The pH values of some common substances are shown in FIGAlthough some very acidic compartments exist within cells (see Chapter 4), most of the interior of an animal or plant cell is neither strongly acidic nor strongly basic but an essentially neutral mixture of acidic and basic substances. Certain bacteria are adapted to life in extremely acidic environments (discussed in Chapter 25), but a substantial change in pH is incompatible with life for most cells. The pH of most types of plant and animal cells (and their environment) ordinarily ranges from around 7.2 to 7.4. URE 2-17.

FIGURE 2-17 Animated pH values of some common solutions A neutral solution (pH 7) has equal concentrations of H+ and OH−. Acidic solutions, which have a higher concentration of H+ than OH−, have pH values less than 7; pH values greater than 7 characterize basic solutions, which have an excess of OH−.

Buffers minimize pH change Many homeostatic mechanisms operate to maintain appropriate pH values. For example, the pH of human blood is about 7.4 and must be maintained within very narrow limits. If the blood becomes too acidic (for example, as a result of respiratory disease), coma and death may result. Excessive alkalinity can result in overexcitability of the nervous system and even convulsions. Organisms contain many natural buffers. A buffer is a substance or combination of substances that resists changes in pH when an acid or base is added. A buffering system includes a weak acid or a weak base. A weak acid or weak base does not ionize completely. At any given instant, only a fraction of the molecules are ionized; most are not dissociated. One of the most common buffering systems functions in the blood of vertebrates (see Chapter 46). Carbon dioxide, produced as a waste product of cell metabolism, enters the blood, the main constituent of which is water. The carbon dioxide reacts with the water to form carbonic acid, a weak acid that dissociates to yield a hydrogen ion and a bicarbonate ion. The following expression describes the buffering system: CO2 + H2 O Carbon dioxide

Water

H2CO3 Carbonic acid

H+ +

HCO 3 – Bicarbonate ion

As the double arrows indicate, all the reactions are reversible. Because carbonic acid is a weak acid, undissociated molecules are always present, as are all the other components of the system. The expression describes the system when it is at dynamic equilibrium, that is, when the rates of the forward and reverse reactions are equal and the relative concentrations of the components are not changing. A system at dynamic equilibrium tends to stay at equilibrium unless a stress is placed on it, which causes it to shift to reduce the stress until it attains a new dynamic equilibrium. A change in the concentration of any component is one such stress. Therefore, the system can be “shifted to the right” by adding reactants or removing products. Conversely, the system can be “shifted to the left” by adding products or removing reactants. Hydrogen ions are the important products to consider in this system. The addition of excess hydrogen ions temporarily shifts the system to the left, as they combine with the bicarbonate ions to form carbonic acid. Eventually, a new dynamic equilibrium is established. At this point the hydrogen ion concentration is similar to the original concentration, and the product of the hydrogen ion and hydroxide ion concentrations is restored to the equilibrium value of 1 × 10−14. If hydroxide ions are added, they combine with the hydrogen ions to form water, effectively removing a product and thus shifting

■ ■

the system to the right. As this occurs, more carbonic acid ionizes, effectively replacing the hydrogen ions that were removed. Organisms contain many weak acids and weak bases, which allows them to maintain an essential reserve of buffering capacity and helps them avoid pH extremes.

An acid and a base react to form a salt When an acid and a base are mixed in water, the H+ of the acid unites with the OH− of the base to form a molecule of water. The remainder of the acid (an anion) combines with the remainder of the base (a cation) to form a salt. For example, hydrochloric acid reacts with sodium hydroxide to form water and sodium chloride: HCl + NaOH ¡ H2O + NaCl

A salt is a compound in which the hydrogen ion of an acid is replaced by some other cation. Sodium chloride, NaCl, is a salt in which the hydrogen ion of HCl has been replaced by the cation Na+. When a salt, an acid, or a base is dissolved in water, its dissociated ions can conduct an electric current; these substances are called electrolytes. Sugars, alcohols, and many other substances do not form ions when dissolved in water; they do not conduct an electric current and are referred to as nonelectrolytes. Cells and extracellular fluids (such as blood) of animals and plants contain a variety of dissolved salts that are the source of the many important mineral ions essential for fluid balance and acid– base balance. Nitrate and ammonium ions from the soil are the important nitrogen sources for plants. In animals, nerve and muscle function, blood clotting, bone formation, and many other aspects of body function depend on ions. Sodium, potassium, calcium, and magnesium are the chief cations present; chloride, bicarbonate, phosphate, and sulfate are important anions. The concentrations and relative amounts of the various cations and anions are kept remarkably constant. Any marked change results in impaired cell functions and may lead to death.

Review ■

■

■

■

A solution has a hydrogen ion concentration of 0.01 mol/L. What is its pH? What is its hydroxide ion concentration? Is it acidic, basic, or neutral? How does this solution differ from one with a pH of 1? What would be the consequences of adding or removing a reactant or a product from a reversible reaction that is at dynamic equilibrium? Why are buffers important in organisms? Why can’t strong acids or bases work as buffers? Why are acids, bases, and salts referred to as electrolytes?

S UM M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

2.1 (page 27) 1 Name the principal chemical elements in living things and provide an important function of each. An element is a substance that cannot be decomposed into simpler substances by normal chemical reactions. About 96% of an organ-

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ism’s mass consists of carbon, the backbone of organic molecules; hydrogen and oxygen, the components of water; and nitrogen, a component of proteins and nucleic acids. 2 Compare the physical properties (mass and charge) and locations of electrons, protons, and neutrons. Distinguish between the atomic number and the mass number of an atom.

Each atom is composed of a nucleus containing positively charged protons and uncharged neutrons. Negatively charged electrons encircle the nucleus. ■ An atom is identified as belonging to a particular element by its number of protons (atomic number). The atomic mass of an atom is equal to the sum of its protons and neutrons. A single proton or a single neutron each has a mass equivalent to ■ one atomic mass unit (amu). The mass of a single electron is only about 1/1800 amu. 3 Define the terms orbital and electron shell. Relate electron shells to principal energy levels. ■ In the space outside the nucleus, electrons move rapidly in electron orbitals. An electron shell consists of electrons in orbitals at the same principal energy level. Electrons in a shell distant from the nucleus have greater energy than those in a shell closer to the nucleus.

Electronegative atoms

■

Learn more about atomic orbitals by clicking on the figure in CengageNOW.

2.2 (page 31) 4 Explain how the number of valence electrons of an atom is related to its chemical properties. The chemical properties of an atom are determined chiefly by the number and arrangement of its most energetic electrons, known as valence electrons. The valence shell of most atoms is full when it contains 8 electrons; that of hydrogen or helium is full when it contains 2. An atom tends to lose, gain, or share electrons to fill its valence shell. 5 Distinguish among simplest, molecular, and structural chemical formulas. ■ Different atoms are joined by chemical bonds to form compounds. A chemical formula gives the types and relative numbers of atoms in a substance. ■ A simplest formula gives the smallest whole-number ratio of the component atoms. A molecular formula gives the actual numbers of each type of atom in a molecule. A structural formula shows the arrangement of the atoms in a molecule. 6 Explain why the mole concept is so useful to chemists. ■ One mole (the atomic or molecular mass in grams) of any substance contains 6.02 × 1023 atoms, molecules, or ions, enabling scientists to “count” particles by weighing a sample. This number is known as Avogadro’s number. ■

2.3 (page 32) 7 Distinguish among covalent bonds, ionic bonds, hydrogen bonds, and van der Waals interactions. Compare them in terms of the mechanisms by which they form and their relative strengths. ■ Covalent bonds are strong, stable bonds formed when atoms share valence electrons, forming molecules. When covalent bonds are formed, the orbitals of the valence electrons may become rearranged in a process known as orbital hybridization. Covalent bonds are nonpolar if the electrons are shared equally between the two atoms. Covalent bonds are polar if one atom is more electronegative (has a greater affinity for electrons) than the other. ■ An ionic bond is formed between a positively charged cation and a negatively charged anion. Ionic bonds are strong in the absence of water but relatively weak in aqueous solution. Learn more about ionic bonding by clicking on the figure in CengageNOW. ■

Hydrogen bonds are relatively weak bonds formed when a hydrogen atom with a partial positive charge is attracted to an atom (usually oxygen or nitrogen) with a partial negative charge already bonded to another molecule or in another part of the same molecule.

O

H

+

–

H N

H

H Hydrogen bond

H

Learn more about hydrogen bonding by clicking on the figure in CengageNOW. ■

van der Waals interactions are weak forces based on fluctuating electric charges.

2.4 (page 36) 8 Distinguish between the terms oxidation and reduction, and relate these processes to the transfer of energy. ■ Oxidation and reduction reactions (redox reactions) are chemical processes in which electrons (and their energy) are transferred from a reducing agent to an oxidizing agent. In oxidation, an atom, ion, or molecule loses electrons (and their energy). In reduction, an atom, ion, or molecule gains electrons (and their energy).

2.5 (page 37) 9 Explain how hydrogen bonds between adjacent water molecules govern many of the properties of water. ■ Water is a polar molecule because one end has a partial positive charge and the other has a partial negative charge. Because its molecules are polar, water is an excellent solvent for ionic or polar solutes. ■ Water molecules exhibit the property of cohesion because they form hydrogen bonds with one another; they also exhibit adhesion through hydrogen bonding to substances with ionic or polar regions. ■ Water has a high heat of vaporization. Hydrogen bonds must be broken for molecules to enter the vapor phase. These molecules carry a great deal of heat, which accounts for evaporative cooling. ■ Because hydrogen bonds must be broken to raise its temperature, water has a high specific heat, which helps organisms maintain a relatively constant internal temperature; this property also helps keep the ocean and other large bodies of water at a constant temperature. ■ The hydrogen bonds between water molecules in ice cause it to be less dense than liquid water. The fact that ice floats makes the aquatic environment less extreme than it would be if ice sank to the bottom. Learn more about the three forms of water by clicking on the figure in CengageNOW.

2.6 (page 40) 10 Contrast acids and bases, and discuss their properties. Acids are proton (hydrogen ion, H+) donors; bases are proton acceptors. An acid dissociates in solution to yield H+ and an anion. Many bases dissociate in solution to yield hydroxide ions (OH−), which then accept protons to form water. 11 Convert the hydrogen ion concentration (moles per liter) of a solution to a pH value and describe how buffers help minimize changes in pH. ■ pH is the negative log of the hydrogen ion concentration of a solution (expressed in moles per liter). A neutral solution with equal concentrations of H+ and OH− (10−7 mol/L) has a pH of 7, an acidic solution has a pH less than 7, and a basic solution has a pH greater than 7. ■

Learn more about the pH of common solutions by clicking on the figure in CengageNOW.

A buffering system is based on a weak acid or a weak base. A buffer resists changes in the pH of a solution when acids or bases are added. 12 Describe the composition of a salt and explain why salts are important in organisms. ■

■

A salt is a compound in which the hydrogen atom of an acid is replaced by some other cation. Salts provide the many mineral ions essential for life functions.

T E S T YO U R U N D E R S TA N D I N G 1. Which of the following elements is mismatched with its properties or function? (a) carbon—forms the backbone of organic compounds (b) nitrogen—component of proteins (c) hydrogen—very electronegative (d) oxygen—can participate in hydrogen bonding (e) all of the above are correctly matched 2.

32 15P,

a radioactive form of phosphorus, has (a) an atomic number of 32 (b) an atomic mass of 15 (c) an atomic mass of 47 (d) 32 electrons (e) 17 neutrons

3. Which of the following facts allows you to determine that atom A and atom B are isotopes of the same element? (a) they each have 6 protons (b) they each have 4 neutrons (c) the sum of the electrons and neutrons in each is 14 (d) they each have 4 valence electrons (e) they each have an atomic mass of 14 4. 11H and 31H have (a) different chemical properties because they have different atomic numbers (b) the same chemical properties because they have the same number of valence electrons (c) different chemical properties because they differ in their number of protons and electrons (d) the same chemical properties because they have the same atomic mass (e) the same chemical properties because they have the same number of protons, electrons, and neutrons 5. The orbitals composing an atom’s valence electron shell (a) are arranged as concentric spheres (b) contain the atom’s least energetic electrons (c) may change shape when covalent bonds are formed (d) never contain more than 1 electron each (e) more than one of the preceding is correct 6. Which of the following bonds and properties are correctly matched? (a) ionic bonds—are strong only if the participating ions are hydrated (b) hydrogen bonds—are responsible for bonding oxygen and hydrogen to form a single water molecule (c) polar covalent bonds—can occur between two atoms of the

same element (d) covalent bonds—may be single, double, or triple (e) hydrogen bonds—are stronger than covalent bonds 7. In a redox reaction (a) energy is transferred from a reducing agent to an oxidizing agent (b) a reducing agent becomes oxidized as it accepts an electron (c) an oxidizing agent accepts a proton (d) a reducing agent donates a proton (e) the electrons in an atom move from its valence shell to a shell closer to its nucleus 8. Water has a high specific heat because (a) hydrogen bonds must be broken to raise its temperature (b) hydrogen bonds must be formed to raise its temperature (c) it is a poor insulator (d) it has low density considering the size of the molecule (e) it can ionize 9. A solution at pH 7 is considered neutral because (a) its hydrogen ion concentration is 0 mol/L (b) its hydroxide ion concentration is 0 mol/L (c) the product of its hydrogen ion concentration and its hydroxide ion concentration is 0 mol/L (d) its hydrogen ion concentration is equal to its hydroxide ion concentration (e) it is nonpolar 10. A solution with a pH of 2 has a hydrogen ion concentration that is _____ the hydrogen ion concentration of a solution with a pH 1 of 4. (a) –12 (b) — 100 (c) 2 times (d) 10 times (e) 100 times 11. Which of the following cannot function as a buffer? (a) phosphoric acid, a weak acid (b) sodium hydroxide, a strong base (c) sodium chloride, a salt that ionizes completely (d) a and c (e) b and c 12. Which of the following statements is true? (a) the number of individual particles (atoms, ions, or molecules) contained in one mole varies depending on the substance (b) Avogadro’s number is the number of particles contained in one mole of a substance (c) Avogadro’s number is 1023 particles (d) one mole of 12C has a mass of 12 g (e) b and d

CRITICAL THINKING 1. Element A has 2 electrons in its valence shell (which is complete when it contains 8 electrons). Would you expect element A to share, donate, or accept electrons? What would you expect of element B, which has 4 valence electrons, and element C, which has 7?

2. A hydrogen bond formed between two water molecules is only –1 as strong as a covalent bond between hydrogen and about 20 oxygen. In what ways would the physical properties of water be different if these hydrogen bonds were stronger (for example, –1 the strength of covalent bonds)? 10 3. Consider the following reaction (in water).

HCl ¡ H+ + Cl− Element A

Element B

Element C

Name the reactant(s) and product(s). Does the expression indicate that the reaction is reversible? Could HCl be used as a buffer?

4. ANALYZING DATA. Could you safely immerse your hand in a solution containing 0.000,000,000,001 g of H+ dissolved in 1 L of water? 5. EVOLUTION LINK. Initiatives designed to discover evidence for life (biosignatures or biomarkers) in the atmospheres of distant planets have been proposed by the U.S. National Aeronautics and Space Administration (NASA) and the European Space Agency (ESA). If implemented, both the Terrestrial Planet Finder project (NASA) and the Darwin project (ESA) will use special space tele-

scopes to detect atmospheric water vapor, as well as oxygen and carbon dioxide. Which of these biosignatures would you consider the most fundamental indicator that life could have evolved on these planets? Why? Additional questions are available in CengageNOW at www.cengage.com/ login.

3

The Chemistry of Life: Organic Compounds

© Eastcott/Momatiuk/The Image Works

Humans obtain organic molecules from food. This young girl is using a leaf to feed her baby brother.

KEY CONCEPTS

B

oth inorganic and organic forms of carbon occur widely in nature. Many types of organic compounds will become incorporated into the

3.1 Carbon atoms join with one another or other atoms to

body of the baby in the photograph as he grows. Organic compounds

form large molecules with a wide variety of shapes. Hydrocarbons are nonpolar, hydrophobic molecules; their properties can be altered by adding functional groups: hydroxyl and carbonyl groups (polar), carboxyl and phosphate groups (acidic), and amino groups (basic).

are those in which carbon atoms are covalently bonded to one another

3.2 Carbohydrates are composed of sugar subunits (mono-

from the breakdown of organic molecules to obtain energy is an example

saccharides), which can be joined to form disaccharides, storage polysaccharides, and structural polysaccharides.

of an inorganic carbon compound. Organic compounds are so named

3.3 Lipids store energy (triacylglycerols) and are the main

living (organic) organisms. In 1828, the German chemist Friedrich Wühler

structural components of cell membranes (phospholipids).

3.4 Proteins have multiple levels of structure and are composed of amino acid subunits joined by peptide bonds.

3.5 Nucleic acids (DNA and RNA) are informational mole-

to form the backbone of the molecule. Some very simple carbon compounds are considered inorganic if the carbon is not bonded to another carbon or to hydrogen. The carbon dioxide we exhale as a waste product

because at one time it was thought that they could be produced only by synthesized urea, a metabolic waste product. Since that time, scientists have learned to synthesize many organic molecules and have discovered organic compounds not found in any organism. Organic compounds are extraordinarily diverse; in fact, more than

cules composed of long chains of nucleotide subunits. ATP and some other nucleotides have a central role in energy metabolism.

5 million have been identified. There are many reasons for this diver-

3.6 Biological molecules have recognizable attributes that

dimensional (3-D) shapes. Furthermore, the carbon atom forms bonds

aid in their identification.

sity. Organic compounds can be produced in a wide variety of three-

with a greater number of different elements than does any other

3.1 CARBON ATOMS AND ORGANIC MOLECULES

type of atom. The addition of chemical groups containing atoms of other elements—especially oxygen, nitrogen, phosphorus, and sulfur—can profoundly change the properties of an organic molecule. Diversity also results from the fact that many organic com-

■ ■ LEARNING OBJECTIVES

pounds found in organisms are extremely large macromolecules,

1

which cells construct from simpler modular subunits. For example,

2

protein molecules are built from smaller compounds called amino acids.

3

As you study this chapter, you will develop an understanding 4

of the major groups of organic compounds found in organisms, including carbohydrates, lipids, proteins, and nucleic acids (DNA

Carbon has unique properties that allow the formation of the carbon backbones of the large, complex molecules essential to life (FIG. 3-1). Because a carbon atom has 4 valence electrons, it can complete its valence shell by forming a total of four covalent bonds (see Fig. 2-2). Each bond can link it to another carbon atom or to an atom of a different element. Carbon is particularly well suited to serve as the backbone of a large molecule because carbon-tocarbon bonds are strong and not easily broken. However, they are not so strong that it would be impossible for cells to break them. Carbon-to-carbon bonds are not limited to single bonds (based

and RNA). Why are these compounds of central importance to all living things? The answer to this question will become more obvious as you study subsequent chapters, in which we explore the evidence that all living things evolved from a common ancestor. Evolution provides a powerful explanation for the similarities of the molecules that constitute the structures of cells and tissues, participate in and regulate metabolic reactions, transmit information, and provide energy for life processes.

H

H

H

C

C

H

H

H H Ethane (a) Chains H H

C

H C

H

H

H

C

C

C

H

Describe the properties of carbon that make it the central component of organic compounds. Define the term isomer and distinguish among the three principal isomer types. Identify the major functional groups present in organic compounds and describe their properties. Explain the relationship between polymers and macromolecules.

H

H H Pr opane

H H H

H

H

C

C

H

H

H H

H

H

C

C

H C

H

H C H

1-Butene

2-Butene

H

H

H

C

C

C

C

H H

H H

C

H H Cyclopentane

C

H

C H

H

H

H Isobutane

H

H

H

H

H

C

C

C

C H

C

H H

H H

H H

H Isopentane

(c) Branched chains

FIGURE 3-1 Organic molecules Note that each carbon atom forms four covalent bonds, producing a wide variety of shapes.

C

C

N

N C

H

H

C

C

H

N H

C

C

H C C

H Benzene

(d) Rings

(b) Double bonds

H

H

C C

H

H

C C

H H

H

O C O H

H

H Histidine (an amino acid) (e) Joined rings and chains

H

H

on sharing one electron pair). Two carbon atoms can share two electron pairs with each other, forming double bonds: C

C

In some compounds, triple carbon-to-carbon bonds are formed: C

C

As shown in Figure 3-1, hydrocarbons, organic compounds consisting only of carbon and hydrogen, can exist as unbranched or branched chains, or as rings. Rings and chains are joined in some compounds. The molecules in the cell are analogous to the components of a machine. Each component has a shape that allows it to fill certain roles and to interact with other components (often with a complementary shape). Similarly, the shape of a molecule is important in determining its biological properties and function. Carbon atoms can link to one another and to other atoms to produce a wide variety of 3-D molecular shapes because the four covalent bonds of carbon do not form in a single plane. Instead, as discussed in Chapter 2, the valence electron orbitals become elongated and project from the carbon atom toward the corners of a tetrahedron (FIG. 3-2). The structure is highly symmetrical, with an angle of about 109.5 degrees between any two of these bonds. Keep in mind that for simplicity, many of the figures in this book are drawn as two-dimensional (2-D) graphic representations of 3-D molecules. Even the simplest hydrocarbon chains, such as those in Figure 3-1, are not actually straight but have a 3-D zigzag structure. Generally, there is freedom of rotation around each carbon-tocarbon single bond. This property permits organic molecules to be flexible and to assume a variety of shapes, depending on the extent to which each single bond is rotated. Double and triple bonds do not allow rotation, so regions of a molecule with such bonds tend to be inflexible.

H

H

C

H H

(b) Methane (CH4)

(a) Carbon (C)

O

C

O

(c) Carbon dioxide (CO2)

FIGURE 3-2 Carbon bonding (a) The 3-D arrangement of the bonds of a carbon atom is responsible for (b) the tetrahedral architecture of methane. (c) In carbon dioxide, oxygen atoms are joined linearly to a central carbon by polar double bonds.

Isomers have the same molecular formula but different structures One reason for the great number of possible carbon-containing compounds is the fact that the same components usually can link in more than one pattern, generating an even wider variety of molecular shapes. Compounds with the same molecular formulas but different structures and thus different properties are called isomers. Isomers do not have identical physical or chemical properties and may have different common names. Cells can distinguish between isomers. Usually, one isomer is biologically active and the other is not. Three types of isomers are structural isomers, geometric isomers, and enantiomers. Structural isomers are compounds that differ in the covalent arrangements of their atoms. For example, FIGURE 3-3a illustrates two structural isomers with the molecular formula C2H6O. Large compounds have more possible structural isomers. There can be up to 366,319 isomers of C20H42. Geometric isomers are compounds that are identical in the arrangement of their covalent bonds but different in the spatial arrangement of atoms or groups of atoms. Geometric isomers are present in some compounds with carbon-to-carbon double bonds. Because double bonds are not flexible, as single bonds are, atoms joined to the carbons of a double bond cannot rotate freely about the axis of the bonds. These cis–trans isomers may be represented as shown in FIGURE 3-3b. The designation cis (Latin, “on this side”) indicates that the two larger components are on the same side of the double bond. If they are on opposite sides of the double bond, the compound is designated a trans (Latin, “across”) isomer. Enantiomers are isomers that are mirror images of each other (FIG. 3-3c). Recall that the four groups bonded to a single carbon atom are arranged at the vertices of a tetrahedron. If the four bonded groups are all different, the central carbon is described as asymmetrical. Figure 3-3c illustrates that the four groups can be arranged around the asymmetrical carbon in two different ways that are mirror images of each other. The two molecules are enantiomers if they cannot be superimposed on each other no matter how they are rotated in space. Although enantiomers have similar chemical properties and most of their physical properties are identical, cells recognize the difference in shape, and usually only one form is found in organisms.

Functional groups change the properties of organic molecules The existence of isomers is not the only source of variety among organic molecules. The addition of various combinations of atoms generates a vast array of molecules with different properties. Because covalent bonds between hydrogen and carbon are nonpolar, hydrocarbons lack distinct charged regions. For this reason, hydrocarbons are insoluble in water and tend to cluster together through hydrophobic interactions. “Water fearing,” the literal meaning of the term hydrophobic, is somewhat misleading. Hydrocarbons interact with water, but much more weakly than the water molecules cohere to one another through hydrogen bonding.

H

H

H

C

C

H

H OH

H

C

O

C

H

H H Ethanol (C2H6O)

H

H

Dimethyl ether (C2H6O)

(a) Structural isomers. Structural isomers differ in the covalent arrangement of their atoms.

H3C

H C

C

H

CH3

H 3C C

CH3

C H

H

trans-2-butene

cis-2-butene

Cengage

(b) Geometric isomers. Geometric, or cis–trans, isomers have identical covalent bonds but differ in the order in which groups are arranged in space.

2

1

C

4

2

3

3

C

1

4

(c) Enantiomers. Enantiomers are isomers that are mirror images of each other. The central carbon is asymmetrical because it is bonded to four different groups. Because of their 3-D structure, the two figures cannot be superimposed no matter how they are rotated.

FIGURE 3-3 Isomers Isomers have the same molecular formula, but their atoms are arranged differently.

or more functional groups, groups of atoms that determine the types of chemical reactions and associations in which the compound participates. Most functional groups readily form associations, such as ionic and hydrogen bonds, with other molecules. Polar and ionic functional groups are hydrophilic because they associate strongly with polar water molecules. The properties of the major classes of biologically important organic compounds—carbohydrates, lipids, proteins, and nucleic acids—are largely a consequence of the types and arrangement of functional groups they contain. When we know what kinds of functional groups are present in an organic compound, we can predict its chemical behavior. Note that the symbol R is used to represent the remainder of the molecule of which each functional group is a part. For example, the methyl group, a common nonpolar hydrocarbon group, is abbreviated ROCH3. As you read the rest of this section, refer to TABLE 3-1 for the structural formulas of other important functional groups, as well as for additional information. The hydroxyl group (abbreviated ROOH) is polar because of the presence of a strongly electronegative oxygen atom. Do not confuse it with the hydroxide ion, OH−, discussed in Chapter 2. If a hydroxyl group replaces one hydrogen of a hydrocarbon, the resulting molecule can have significantly altered properties. For example, ethane (see Fig. 3-1a) is a hydrocarbon that is a gas at room temperature. If a hydroxyl group replaces a hydrogen atom, the resulting molecule is ethyl alcohol, or ethanol, which is found in alcoholic beverages (see Fig. 3-3a). Ethanol is somewhat cohesive because the polar hydroxyl groups of adjacent molecules interact; it is therefore liquid at room temperature. Unlike ethane, ethyl alcohol dissolves in water because the polar hydroxyl groups interact with the polar water molecules. The carbonyl group consists of a carbon atom that has a double covalent bond with an oxygen atom. This double bond is polar because of the electronegativity of the oxygen; thus, the carbonyl group is hydrophilic. The position of the carbonyl group in the molecule determines the class to which the molecule belongs. An aldehyde has a carbonyl group positioned at the end of the carbon skeleton (abbreviated ROCHO); a ketone has an internal carbonyl group (abbreviated ROCOOR). The carboxyl group (abbreviated ROCOOH) in its nonionized form consists of a carbon atom joined by a double covalent bond to an oxygen atom, and by a single covalent bond to another oxygen, which is in turn bonded to a hydrogen atom. Two electronegative oxygen atoms in such close proximity establish an extremely polarized condition, which can cause the hydrogen atom to be stripped of its electron and released as a hydrogen ion (H+). The resulting ionized carboxyl group has 1 unit of negative charge (ROCOO−): O

O

Hydrocarbons interact weakly with one another, but the main reason for hydrophobic interactions is that they are driven together in a sense, having been excluded by the hydrogen-bonded water molecules. However, the characteristics of an organic molecule can be changed dramatically by replacing one of the hydrogens with one

R

¡ R

C

+ H+

C –

O

O H

Carboxyl groups are weakly acidic; only a fraction of the molecules ionize in this way. This group therefore exists in one of

TABLE 3-1

Some Biologically Important Functional Groups

Functional Group and Description

Structural Formula

Class of Compound Characterized by Group

Hydroxyl Polar because electronegative oxygen attracts covalent electrons

R

Alcohols

OH

H

H

H

C

C

H

H

OH

Example, ethanol Carbonyl Aldehydes: Carbonyl group carbon is bonded to at least one H atom; polar because electronegative oxygen attracts covalent electrons Ketones: Carbonyl group carbon is bonded to two other carbons; polar because electronegative oxygen attracts covalent electrons

Aldehydes

O R

C

O

H

H

C

H

Example, formaldehyde Ketones

O R

C

R

H

H

O

H

C

C

C

H

H

H

Example, acetone Carboxyl Weakly acidic; can release an H+

O R

C

Carboxylic acids (organic acids)

O R

OH

Non-ionized

C

O– + H+

Ionized

O R

C

OH

Example, amino acid Amino Weakly basic; can accept an H+

R

N

H

R

H

Non-ionized

N+

Amines

H H H

NH2 O R

Ionized

C

C

OH

H Example, amino acid Phosphate Weakly acidic; one or two H+ can be released

O R

O

P

OH

R

O

P O–

OH Non-ionized

Organic Phosphates

O O–

O HO

P

O

R

OH

Ionized

Example, phosphate ester (as found in ATP) Sulfhydryl Helps stabilize internal structure of proteins

R

SH

Thiols

H

H

H

O

C

C

C

OH

SH NH2 Example, cysteine

two hydrophilic states: ionic or polar. Carboxyl groups are essential constituents of amino acids. An amino group (abbreviated RONH2) in its non-ionized form includes a nitrogen atom covalently bonded to two hydrogen atoms. Amino groups are weakly basic because they are able to ac-

cept a hydrogen ion (proton). The resulting ionized amino group has 1 unit of positive charge (RONH3+). Amino groups are components of amino acids and of nucleic acids. A phosphate group (abbreviated ROPO4H2) is weakly acidic. The attraction of electrons by the oxygen atoms can result in

Monomer

FIGURE 3-4 A simple polymer This small polymer of polyethylene is formed by linking two-carbon ethylene (C2H4) monomers. One such monomer is outlined in red. The structure is represented by a space-filling model, which accurately depicts the 3-D shape of the molecule.

lated by a specific enzyme (biological catalyst), a hydrogen from a water molecule attaches to one monomer, and a hydroxyl from water attaches to the adjacent monomer (FIG. 3-5). Monomers become covalently linked by condensation reactions. Because the equivalent of a molecule of water is removed during the reactions that combine monomers, the term dehydration synthesis is sometimes used to describe condensation (see Fig. 3-5). However, in biological systems the synthesis of a polymer is not simply the reverse of hydrolysis, even though the net effect is the opposite of hydrolysis. Synthetic processes such as condensation require energy and are regulated by different enzymes. In the following sections we examine carbohydrates, lipids, proteins, and nucleic acids. Our discussion begins with the smaller, simpler forms of these compounds and extends to the linking of these monomers to form macromolecules.

Review ■

the release of one or two hydrogen ions, producing ionized forms with 1 or 2 units of negative charge. Phosphates are constituents of nucleic acids and certain lipids. The sulfhydryl group (abbreviated ROSH), consisting of an atom of sulfur covalently bonded to a hydrogen atom, is found in molecules called thiols. As you will see, amino acids that contain a sulfhydryl group can make important contributions to the structure of proteins.

■

■

■

■

Many biological molecules are polymers Many biological molecules such as proteins and nucleic acids are very large, consisting of thousands of atoms. Such giant molecules are known as macromolecules. Most macromolecules are polymers, produced by linking small organic compounds called monomers (FIG. 3-4). Just as all the words in this book have been written by arranging the 26 letters of the alphabet in various combinations, monomers can be grouped to form an almost infinite variety of larger molecules. The thousands of different complex organic compounds present in organisms are constructed from about 40 small, simple monomers. For example, the 20 monomers called amino acids can be linked end to end in countless ways to form the polymers known as proteins. Polymers can be degraded to their component monomers by hydrolysis reactions (“to break with water”). In a reaction regu-

What are some of the ways that the features of carbon-to-carbon bonds influence the stability and 3-D structure of organic molecules? Draw pairs of simple sketches comparing two (1) structural isomers, (2) geometric isomers, and (3) enantiomers. Why are these differences biologically important? Sketch the following functional groups: methyl, amino, carbonyl, hydroxyl, carboxyl, and phosphate. Include both non-ionized and ionized forms for acidic and basic groups. How is the fact that a group is nonpolar, polar, acidic, or basic related to its hydrophilic or hydrophobic properties? Why is the equivalent of a water molecule important to both condensation reactions and hydrolysis reactions?

3.2 CARBOHYDRATES ■ ■ LEARNING OBJECTIVE 5

Distinguish among monosaccharides, disaccharides, and polysaccharides; compare storage polysaccharides with structural polysaccharides.

Sugars, starches, and cellulose are carbohydrates. Sugars and starches serve as energy sources for cells; cellulose is the main structural component of the walls that surround plant cells. Carbohydrates contain carbon, hydrogen, and oxygen atoms in a ratio of approximately one carbon to two hydrogens to one oxygen (CH2O)n. The term carbohydrate, meaning “hydrate (water) of

Condensation Enzyme A HO

OH Monomer

HO

OH Monomer

HO Hydrolysis Enzyme B

FIGURE 3-5 Animated Condensation and hydrolysis reactions Joining two monomers yields a dimer; incorporating additional monomers produces a polymer. Note that condensation and hydrolysis reactions are catalyzed by different enzymes.

O Dimer

OH +

H2O

The large number of polar hydroxyl groups, plus the carbonyl group, gives a monosaccharide hydrophilic properties. FIGURE 3-6 shows simplified, 2-D representations of some common monosaccharides. The simplest carbohydrates are the three-carbon sugars (trioses): glyceraldehyde and dihydroxyacetone. Ribose and deoxyribose are common pentoses, sugars that contain five carbons; they are components of nucleic acids (DNA, RNA, and related compounds). Glucose, fructose, galactose, and other six-carbon sugars are called hexoses. (Note that the names of carbohydrates typically end in -ose.) Glucose (C6H12O6), the most abundant monosaccharide, is used as an energy source in most organisms. During cellular respiration (see Chapter 8), cells oxidize glucose molecules, converting the stored energy to a form that can be readily used for cell work. Glucose is also used in the synthesis of other types of compounds

carbon,” reflects the 2:1 ratio of hydrogen to oxygen, the same ratio found in water (H2O). Carbohydrates contain one sugar unit (monosaccharides), two sugar units (disaccharides), or many sugar units (polysaccharides).

Monosaccharides are simple sugars Monosaccharides typically contain from three to seven carbon atoms. In a monosaccharide, a hydroxyl group is bonded to each carbon except one; that carbon is double-bonded to an oxygen atom, forming a carbonyl group. If the carbonyl group is at the end of the chain, the monosaccharide is an aldehyde; if the carbonyl group is at any other position, the monosaccharide is a ketone. (By convention, the numbering of the carbon skeleton of a sugar begins with the carbon at or nearest the carbonyl end of the open chain.)

H H

H O

H

1

C 2

C

H

C

2

C

OH

3

C

H

1

H

3

C

H

OH

OH

H

O

H

OH

H

1

C

2

C

3

C

4

C

5

C

O

H

OH

H

OH

H

OH

H

OH

H

1

C

2

C

3

C

4

C

5

C

O H OH OH OH

H

H

H

H

Glyceraldehyde (C3H6O3) (an aldehyde)

Dihydroxyacetone (C3H6O3) (a ketone)

Ribose (C5H10O5) (the sugar component of RNA)

Deoxyribose (C5H10O4) (the sugar component of DNA)

(a) Triose sugars (3-carbon sugars)

(b) Pentose sugars (5-carbon sugars)

H H H HO H H H

1

C

2

C

3

C

4

C

5

C

6

C

O

H

C

2

OH H

1

C

HO

OH

H

OH

H

OH

H

3

C

4

C

5

C

6

C

OH

H

O

H

H

HO

OH

HO

OH

H

OH

H

1

C

2

C

3

C

4

C

5

C

6

C

O OH H H OH OH

H

H

H

Glucose (C6H12O6) (an aldehyde)

Fructose (C6H12O6) (a ketone)

Galactose (C6H12O6) (an aldehyde)

(c) Hexose sugars (6-carbon sugars)

FIGURE 3-6 Monosaccharides Shown are 2-D chain structures of (a) three-carbon trioses, (b) five-carbon pentoses, and (c) six-carbon hexoses. Although it is convenient to show monosaccharides in this form, the pentoses and hexoses are more accurately depicted as ring structures, as in Figure 3-7. The carbonyl group (gray screen) is terminal in aldehyde sugars and located in an internal position in ketones. Deoxyribose differs from ribose because deoxyribose has one less oxygen; a hydrogen (white screen) instead of a hydroxyl group (blue screen) is attached to carbon 2. Glucose and galactose differ in the arrangement of the hydroxyl group and hydrogen attached to carbon 4 (red box).

such as amino acids and fatty acids. GluOH OH OH cose is so important in metabolism that 6 6 6 mechanisms have evolved to maintain H C H H C H H C H 5 5 5 its concentration at relatively constant C C C O H OH O O OH H H H levels in the blood of humans and other 4 4 4 H H 1 1 1 OH H C H C H C H C C C complex animals (see Chapter 49). OH OH 3 2 3 2 3 2 Glucose and fructose are structural HO C HO C HO C C OH C C H isomers: they have identical molecular H OH H OH H OH formulas, but their atoms are arranged differently. In fructose (a ketone) the 𝛂-Glucose 𝛃-Glucose double-bonded oxygen is linked to a Formation of glucose ring (ring form) (ring form) carbon within the chain rather than to a (a) When dissolved in water, glucose undergoes a rearrangement of its atoms, forming one of two possible ring structures: α-glucose or β-glucose. Although terminal carbon as in glucose (an aldethe drawing does not show the complete 3-D structure, the thick, tapered bonds hyde). Because of these differences, the in the lower portion of each ring represent the part of the molecule that would two sugars have different properties. For project out of the page toward you. example, fructose, found in honey and some fruits, tastes sweeter than glucose. CH2OH CH2OH Glucose and galactose are both hexO O OH oses and aldehydes. However, they differ OH OH in the arrangement of the atoms attached OH HO HO to asymmetrical carbon atom 4. OH OH The linear formulas in Figure 3-6 give a clear but somewhat unrealistic 𝛂-Glucose 𝛃-Glucose picture of the structures of some com(b) The essential differences between α-glucose and β-glucose are more readily mon monosaccharides. As we have menapparent in these simplified structures. By convention, a carbon atom is assumed to be present at each angle in the ring unless another atom is shown. Most tioned, molecules are not 2-D; in fact, hydrogen atoms have been omitted. the properties of each compound depend largely on its 3-D structure. Thus, 3-D formulas are helpful in understandFIGURE 3-7 a and b forms of glucose ing the relationship between molecular structure and biological function. Molecules of glucose and other pencarbon 1 of one molecule and carbon 4 of the other molecule. toses and hexoses in solution are actually rings rather than extended The disaccharide maltose (malt sugar) consists of two covalently straight carbon chains. linked a-glucose units. Sucrose, common table sugar, consists of Glucose in solution (as in the cell) typically exists as a ring of a glucose unit combined with a fructose unit. Lactose (the sugar five carbons and one oxygen. It assumes this configuration when present in milk) consists of one molecule of glucose and one of its atoms undergo a rearrangement, permitting a covalent bond to galactose. connect carbon 1 to the oxygen attached to carbon 5 (FIG. 3-7). When glucose forms a ring, two isomeric forms are possible, difAs shown in Figure 3-8, a disaccharide can be hydrolyzed, that fering only in orientation of the hydroxyl (OOH) group attached is, split by the addition of water, into two monosaccharide units. to carbon 1. When this hydroxyl group is on the same side of the During digestion, maltose is hydrolyzed to form two molecules of plane of the ring as the OCH2OH side group, the glucose is desglucose: ignated beta glucose (b-glucose). When it is on the side (with remaltose + water ¡ glucose + glucose spect to the plane of the ring) opposite the OCH2OH side group, the compound is designated alpha glucose (a-glucose). Although Similarly, sucrose is hydrolyzed to form glucose and fructose: the differences between these isomers may seem small, they have important consequences when the rings join to form polymers. sucrose + water ¡ glucose + fructose

Disaccharides consist of two monosaccharide units A disaccharide (two sugars) contains two monosaccharide rings joined by a glycosidic linkage, consisting of a central oxygen covalently bonded to two carbons, one in each ring (FIG. 3-8). The glycosidic linkage of a disaccharide generally forms between

Polysaccharides can store energy or provide structure A polysaccharide is a macromolecule consisting of repeating units of simple sugars, usually glucose. The polysaccharides are the most abundant carbohydrates and include starches, glycogen, and

6

H 4

HO

Glycosidic linkage 6 CH2OH

CH2OH 5

H OH 3

H

O H

H

H 1

4

2

O

5

O

OH

H

H

H OH

H

1

3

2

OH

H

CH2OH

CH2OH

+

H2O

O H OH

Enzyme

OH

OH H

Maltose C12H22O11

O H OH

+

H

HO

H

H

H

H

HO

OH

OH

H

OH

Glucose C6H12O6

Glucose C6H12O6

(a) Maltose may be broken down (as during digestion) to form two molecules of glucose.The glycosidic linkage is broken in a hydrolysis reaction, which requires the addition of water. CH2OH H

CH2OH O

H OH

H

HOCH2

H

HO

O H

HO

O H

OH

H

H +

H2O

OH

H OH

Enzyme

CH2OH H

Sucrose C12H22O11

O

H

+

H

HO

OH H

OH

Glucose C6 H12O6

1

HO CH2 2

HO

O H 3

OH

H HO

5

4

6

CH2OH

H

Fructose C6 H12O6

(b) Sucrose can be hydrolyzed to yield a molecule of glucose and a molecule of fructose.

FIGURE 3-8 Hydrolysis of disaccharides Note that an enzyme is needed to promote these reactions.

cellulose. Although the precise number of sugar units varies, thousands of units are typically present in a single molecule. A polysaccharide may be a single long chain or a branched chain. Because they are composed of different isomers and because the units may be arranged differently, polysaccharides vary in their properties. Those that can be easily broken down to their subunits are well suited for energy storage, whereas the macromolecular 3-D architecture of others makes them particularly well suited to form stable structures. Starch, the typical form of carbohydrate used for energy storage in plants, is a polymer consisting of a-glucose subunits. These monomers are joined by a 1O4 linkages, which means that carbon 1 of one glucose is linked to carbon 4 of the next glucose in the chain (FIG. 3-9). Starch occurs in two forms: amylose and amylopectin. Amylose, the simpler form, is unbranched. Amylopectin, the more common form, usually consists of about 1000 glucose units in a branched chain. Plant cells store starch mainly as granules within specialized organelles called amyloplasts (see Fig. 3-9a); some cells, such as those of potatoes, are very rich in amyloplasts. When energy is needed for cell work, the plant hydrolyzes the starch, releasing the glucose subunits. Virtually all organisms, including humans and other animals, have enzymes that can break a 1O4 linkages.

Glycogen (sometimes referred to as animal starch) is the form in which glucose subunits, joined by a 1O4 linkages, are stored as an energy source in animal tissues. Glycogen is similar in structure to plant starch but more extensively branched and more water soluble. In vertebrates, glycogen is stored mainly in liver and muscle cells. Carbohydrates are the most abundant group of organic compounds on Earth, and cellulose is the most abundant carbohydrate; it accounts for 50% or more of all the carbon in plants (FIG. 3-10). Cellulose is a structural carbohydrate. Wood is about half cellulose, and cotton is at least 90% cellulose. Plant cells are surrounded by strong supporting cell walls consisting mainly of cellulose. Cellulose is an insoluble polysaccharide composed of many joined glucose molecules. The bonds joining these sugar units are different from those in starch. Recall that starch is composed of aglucose subunits, joined by a 1O4 glycosidic linkages. Cellulose contains b-glucose monomers joined by b 1O4 linkages. These bonds cannot be split by the enzymes that hydrolyze the a linkages in starch. Because humans, like other animals, lack enzymes that digest cellulose, we cannot use it as a nutrient. The cellulose found in whole grains and vegetables remains fibrous and provides bulk that helps keep our digestive tract functioning properly. Some microorganisms digest cellulose to glucose. In fact, cellulose-digesting bacteria live in the digestive systems of cows

Ed Reschke

and sheep, enabling these grass-eating animals to obtain nourishment from cellulose. Similarly, the digestive systems of termites contain microorganisms that digest cellulose (see Fig. 26-4b). Cellulose molecules are well suited for a structural role. The bglucose subunits are joined in a way that allows extensive hydrogen bonding among different cellulose molecules, and they aggregate in long bundles of fibers (see Fig. 3-10a).

Some modified and complex carbohydrates have special roles

Many derivatives of monosaccharides are important biological molecules. Some form important structural components. The amino sugars galactosamine and glucosamine are compounds in which a hydroxyl group (OOH) is replaced by an amino group (ONH2). Galactosamine is present in cartilage, a constituent of the skeletal system of vertebrates. N-acetyl glucosamine (NAG) subunits, joined by glycosidic bonds, Amyloplasts compose chitin, a main component of the cell walls of fungi and of the external skeletons of insects, crayfish, and other arthropods (FIG. 3-11). Chitin forms very tough structures because, as in cellulose, its molecules interact through multiple hydrogen bonds. Some chitinous structures, such as the shell of a lobster, are further hardened by the addition of calcium carbonate (CaCO3), an inorganic form of carbon. Carbohydrates may also combine with proteins to form glycoproteins, compounds present on the outer surface of cells other than bacteria. Some of these carbohydrate chains allow cells to adhere to one another, whereas others provide protection. Most proteins secreted by cells are glycoproteins. These include the major components of mucus, a protective material secreted by the mucous membranes of the respiratory and digestive systems. Carbohydrates combine with lipids to form glycolipids, com100 μm pounds on the surfaces of animal cells that allow cells to (a) Starch (stained purple) is stored in specialized recognize and interact with one another. organelles, called amyloplasts, in these cells of a buttercup root.

6

CH2OH

O

H OH H

O H H 4

H

O

OH

CH2OH 5

O H

3

2

H OH H

H

HO O 6 CH 2

CH2OH H HO

H OH H

1

O H H

O H H H OH

O

CH2OH

OH

H

H

OH

O

CH2OH O H H

OH

H

H

OH

O

H OH H

O H H

O

OH

(b) Starch is composed of α-glucose molecules joined by glycosidic bonds. At the branch points are bonds between carbon 6 of the glucose in the straight chain and carbon 1 of the glucose in the branching chain. (c) Starch consists of highly branched chains; the arrows indicate the branch points. Each chain is actually a coil or helix, stabilized by hydrogen bonds between the hydroxyl groups of the glucose subunits.

FIGURE 3-9 Starch, a storage polysaccharide

Review ■

■

3.3 LIPIDS

What features related to hydrogen bonding give storage polysaccharides, such as starch and glycogen, different properties from structural polysaccharides, such as cellulose and chitin? Why can’t humans digest cellulose?

■ ■ LEARNING OBJECTIVE 6 Distinguish among fats, phospholipids, and steroids, and describe the composition, characteristics, and biological functions of each.

Biophoto Associates/ Photo Researchers, Inc.

Unlike carbohydrates, which are defined by their structure, lipids are a heterogeneous group of compounds that are categorized by the fact that they are soluble in nonpolar solvents (such as ether and chloroform) and are relatively insoluble in water. Lipid molecules have these properties because they consist mainly of carbon and hydrogen, with few oxygencontaining functional groups. Hydrophilic functional groups typically contain oxygen atoms; therefore, lipids, which have little oxygen, tend to be hydrophobic. Among the biologically important groups of lipids are fats, phospholipids, carotenoids (orange and yellow plant pigments), steroids, and waxes. Some lipids are used for energy storage, others serve as structural components of cell membranes, and some are important hormones.

1 μm

(a) Cellulose fibers from a cell wall. The fibers shown in this electron micrograph consist of bundles of cellulose molecules that interact through hydrogen bonds.

CH2OH H HO

O H OH

O

H H

H

H

OH

OH H

H

H

OH

CH2OH H

H O

O CH2OH

O H OH

O

H H

H

H

OH

OH H

H

H

H

O

O

CH2OH

OH

(b) The cellulose molecule is an unbranched polysaccharide. It consists of about 10,000 ß-glucose units joined by glycosidic bonds.

FIGURE 3-10 Cellulose, a structural polysaccharide

N-acetyl glucosamine

H

H H

H

NHCOCH3

OH H

H

O

O H OH

H

H

NHCOCH3

O

H

H

CH2OH

CH2OH

H H

O H

OH H

H

H

NHCOCH3

(a) Chitin is a polymer composed of N-acetyl glucosamine subunits.

FIGURE 3-11 Chitin, a structural polysaccharide

NHCOCH3

O

O H OH

H

O CH2OH

H O

Dwight R. Kuhn

CH2OH

(b) Chitin is an important component of the exoskeleton (outer covering) this dragonfly is shedding.

Triacylglycerol is formed from glycerol and three fatty acids

Saturated and unsaturated fatty acids differ in physical properties

The most abundant lipids in living organisms are triacylglycerols. These compounds, commonly known as fats, are an economical form of reserve fuel storage because, when metabolized, they yield more than twice as much energy per gram as do carbohydrates. Carbohydrates and proteins can be transformed by enzymes into fats and stored within the cells of adipose (fat) tissue of animals and in some seeds and fruits of plants. A triacylglycerol molecule (also known as a triglyceride) consists of glycerol joined to three fatty acids (FIG. 3-12). Glycerol is a three-carbon alcohol that contains three hydroxyl (OOH) groups, and a fatty acid is a long, unbranched hydrocarbon chain with a carboxyl group (OCOOH) at one end. A triacylglycerol molecule is formed by a series of three condensation reactions. In each reaction, the equivalent of a water molecule is removed as one of the glycerol’s hydroxyl groups reacts with the carboxyl group of a fatty acid, resulting in the formation of a covalent linkage known as an ester linkage (see Fig. 3-12b). The first reaction yields a monoacylglycerol (monoglyceride); the second, a diacylglycerol (diglyceride); and the third, a triacylglycerol. During digestion, triacylglycerols are hydrolyzed to produce fatty acids and glycerol (see Chapter 47). Diacylglycerol is an important molecule for sending signals within the cell (see Chapters 6 and 49).

About 30 different fatty acids are commonly found in lipids, and they typically have an even number of carbon atoms. For example, butyric acid, present in rancid butter, has four carbon atoms. Oleic acid, with 18 carbons, is the most widely distributed fatty acid in nature and is found in most animal and plant fats. Saturated fatty acids contain the maximum possible number of hydrogen atoms. Palmitic acid, a 16-carbon fatty acid, is a common saturated fatty acid (see Fig. 3-12c). Fats high in saturated fatty acids, such as animal fat and solid vegetable shortening, tend to be solid at room temperature. The reason is that even electrically neutral, nonpolar molecules can develop transient regions of weak positive charge and weak negative charge. This occurs as the constant motion of their electrons causes some regions to have a temporary excess of electrons, whereas others have a temporary electron deficit. These slight opposite charges result in van der Waals interactions between adjacent molecules (see Chapter 2). Although van der Waals interactions are weak attractions, they are strong when many occur among long hydrocarbon chains. These van der Waals interactions tend to make a substance more solid by limiting the motion of its molecules. Unsaturated fatty acids include one or more adjacent pairs of carbon atoms joined by a double bond. Therefore, they are not fully saturated with hydrogen. Fatty acids with one double bond are monounsaturated fatty acids, whereas those with more than one double bond are polyunsaturated fatty acids. Oleic acid is a monounsaturated fatty acid, and linoleic acid is a common polyunsaturated fatty acid (see Figs. 3-12d and e). Fats containing a high proportion of monounsaturated or polyunsaturated fatty acids tend to be liquid at room temperature. The reason is that each double bond produces a bend in the hydrocarbon chain that prevents it from aligning closely with an adjacent chain, thereby limiting van der Waals interactions and permitting freer molecular motion.

H H

C

OH

H

C

OH

H

C

OH

Carboxyl O HO

H Glycerol

(a) H H

H

H

C

C

C H

(b)

C

R

Fatty acid

O

O

O

Ester linkage O H H

H

H

H

H

H

H

H

H

H

H

H

H

C

C

C

C

C

C

C

C

C

C

C

C

C

C

C

H

H

H

H

H

H

H

H

H

H

H

H

H

H

O

H

H

H

H

H

H

H

H

H

H

H

H

H

H

H

H

C

C

C

C

C

C

C

C

C

C

C

C

C

C

C

C

C

H

H

H

H

H

H

H

H

H

H

H

H

H

H

O

H

H

H

H

H

H

H

H

H

H

H

H

H

H

H

H

C

C

C

C

C

C

C

C

C

C

C

C

C

C

C

C

C

H

H

H

H

H

H

H

H

H

H

H

H

A triacylglycerol

FIGURE 3-12 Animated Triacylglycerol, the main storage lipid (a) Glycerol and fatty acids are the components of fats. (b) Glycerol is attached to fatty acids by ester linkages (in gray). The space-filling models show the actual shapes of the fatty acids. (c) Palmitic acid, a saturated fatty acid, is a straight chain. (d) Oleic acid (monounsaturated) and (e) linoleic acid (polyunsaturated) are bent or kinked wherever a carbon-to-carbon double bond appears.

CH3

(c) Palmitic acid CH3

(d) Oleic acid CH3

(e) Linoleic acid

does not produce a bend at the site of the double bond, trans fatty acids are more solid at room temperature; like saturated fatty acids, they increase the risk of cardiovascular disease. At least two unsaturated fatty acids (linoleic acid and arachidonic acid) are essential nutrients that must be obtained from food because the human body cannot synthesize them. However, the amounts required are small, and deficiencies are rarely seen. There is no dietary requirement for saturated fatty acids.

Food manufacturers commonly hydrogenate or partially hydrogenate cooking oils to make margarine and other foodstuffs, converting unsaturated fatty acids to saturated fatty acids and making the fat more solid at room temperature. This process makes the fat less healthful because saturated fatty acids in the diet are known to increase the risk of cardiovascular disease (see Chapter 44). The hydrogenation process has yet another effect. Note that in the naturally occurring unsaturated fatty acids oleic acid and linoleic acid shown in Figure 3-12, the two hydrogens flanking each double bond are on the same side of the hydrocarbon chain (the cis configuration). When fatty acids are artificially hydrogenated, the double bonds can become rearranged, resulting in a trans configuration, analogous to the arrangement shown in Figure 3-3b. Trans fatty acids are technically unsaturated, but they mimic many of the properties of saturated fatty acids. Because the trans configuration

KEY POINT

N+

Phospholipids belong to a group of lipids called amphipathic lipids, in which one end of each molecule is hydrophilic and the other end is hydrophobic (FIG. 3-13). The two ends of a phospho-

A lipid bilayer forms when phospholipids interact with water.

O

CH3 CH3

Phospholipids are components of cell membranes

CH2

CH2

O

P

H O

C

H O

O–

CH3

H

H

C

C H

Choline

O

O

C

H C

H C

H C

H C

H C

H C

H

H C

C

C

H

H

H

H

H

CH 3

C

C

C

C

C

C

C

H

H

H

H

H

H

H

H

H

H

H

H

H

H

H

H

H

O

H

H

H

H

H

H

H

H

H

H

H

H

H

H

H

H

C

C

C

C

C

C

C

C

C

C

C

C

C

C

C

C

C

H

H

H

H

H

H

H

H

H

H

H

H

H

H

H

H

Phosphate Glycerol group

CH3

Fatty acids

Water

Hydrophilic head

Hydrophobic tail

(a) Phospholipid (lecithin). A phospholipid consists of a hydrophobic tail, made up of two fatty acids, and a hydrophilic head, which includes a glycerol bonded to a phosphate group, which is in turn bonded to an organic group that can vary. Choline is the organic group in lecithin (or phosphatidylcholine), the molecule shown. The fatty acid at the top of the figure is monounsaturated; it contains one double bond that produces a characteristic bend in the chain.

FIGURE 3-13 Animated A phospholipid and a phospholipid bilayer

(b) Phospholipid bilayer. Phospholipids form lipid bilayers in which the hydrophilic heads interact with water and the hydrophobic tails are in the bilayer interior.

lipid differ both physically and chemically. A phospholipid consists of a glycerol molecule attached at one end to two fatty acids and at the other end to a phosphate group linked to an organic compound such as choline. The organic compound usually contains nitrogen. (Note that phosphorus and nitrogen are absent in triacylglycerols, as shown in Figure 3-12b.) The fatty acid portion of the molecule (containing the two hydrocarbon “tails”) is hydrophobic and not soluble in water. However, the portion composed of glycerol, phosphate, and the organic base (the “head” of the molecule) is ionized and readily water soluble. The amphipathic properties of phospholipids cause them to form lipid bilayers in aqueous (watery) solution. Thus, they are uniquely suited as the fundamental components of cell membranes (discussed in Chapter 5).

CH2

CH2 CH2 CH3 C

C C

CH3

CH2

CH2 CH3

C

C

C C

CH3

CH C

CH3

CH CH3

C

HC

CH

CH HC

HC

(a) Isoprene

C

C

CH3

The orange and yellow plant pigments called carotenoids are classified with the lipids because they are insoluble in water and have an oily consistency. These pigments, found in the cells of plants, play a role in photosynthesis. Carotenoid molecules, such as b-carotene, and many other important pigments, consist of five-carbon hydrocarbon monomers known as isoprene units (FIG. 3-14). Most animals convert carotenoids to vitamin A, which can then be converted to the visual pigment retinal. Three groups of animals—the mollusks, insects, and vertebrates—have eyes that use retinal in the process of light reception. Notice that carotenoids, vitamin A, and retinal all have a pattern of double bonds alternating with single bonds. The electrons that make up these bonds can move about relatively easily when light strikes the molecule. Such molecules are pigments; they tend to be highly colored because the mobile electrons cause them to strongly absorb light of certain wavelengths and reflect light of other wavelengths.

H

CH

Point of cleavage

HC

CH3 CH2

CH

CH2 C

CH3

C C

CH3

HC

CH3

HC

CH

CH3

CH2

CH3

HC

CH

C

CH3

OH

H

HC

C

C

(c) Vitamin A

HC C

CH3

HC

HC

Carotenoids and many other pigments are derived from isoprene units

CH3

HC

CH

CH2

CH3

HC

HC CH2

CH2

CH3

C

C

CH2

CH2

C CH3

CH3

HC CH

CH2

HC

(b) 𝛃-Carotene

C

CH3

HC C

Steroids contain four rings of carbon atoms A steroid consists of carbon atoms arranged in four attached rings; three of the rings contain six carbon atoms, and the fourth contains five (FIG. 3-15). The length and structure of the side chains that extend from these rings distinguish one steroid from another. Like carotenoids, steroids are synthesized from isoprene units. Among the steroids of biological importance are cholesterol, bile salts, reproductive hormones, and cortisol as well as other hormones secreted by the adrenal cortex. Cholesterol is an essential structural component of animal cell membranes, but when excess cholesterol in blood forms plaques on artery walls, the risk of cardiovascular disease increases (see Chapter 44). Plant cell membranes contain molecules similar to cholesterol. Interestingly, some of these plant steroids block the intestine’s absorption of cholesterol. Bile salts emulsify fats in the intestine so they can be enzymatically hydrolyzed. Steroid hormones regulate certain aspects of metabolism in a variety of animals and plants.

H

O

(d) Retinal

FIGURE 3-14 Isoprene-derived compounds (a) An isoprene subunit. (b) b-carotene, with dashed lines indicating the boundaries of the individual isoprene units within. The wavy line is the point at which most animals cleave the molecule to yield two molecules of (c) vitamin A. Vitamin A is converted to the visual pigment (d) retinal.

Some chemical mediators are lipids Animal cells secrete chemicals to communicate with one another or to regulate their own activities. Some chemical mediators are produced by the modification of fatty acids that have been removed from membrane phospholipids. These include prostaglandins, which have varied roles, including promoting inflammation and smooth muscle contraction. Certain hormones, such as

CH3

CH2

CH

CH2

CH2

CH3 CH

CH3

CH3

CH3

TABLE 3-2

Major Classes of Proteins and Their Functions

Protein Class

Functions and Examples

Enzymes

Catalyze specific chemical reactions

Structural proteins

Strengthen and protect cells and tissues (e.g., collagen strengthens animal tissues)

Storage proteins

Store nutrients; particularly abundant in eggs (e.g., ovalbumin in egg white) and seeds (e.g., zein in corn kernels)

Transport proteins

Transport specific substances between cells (e.g., hemoglobin transports oxygen in red blood cells); move specific substances (e.g., ions, glucose, amino acids) across cell membranes

Regulatory proteins

Some are protein hormones (e.g., insulin); some control the expression of specific genes

Motile proteins

Participate in cellular movements (e.g., actin and myosin are essential for muscle contraction)

Protective proteins

Defend against foreign invaders (e.g., antibodies play a role in the immune system)

HO Indicates double bond

(a) Cholesterol is an essential component of animal cell membranes. CH2OH

HO

CH3

C

O OH

CH3

O

(b) Cortisol is a steroid hormone secreted by the adrenal glands.

■ ■ LEARNING OBJECTIVES

Proteins, macromolecules composed of amino acids, are the most versatile cell components. As will be discussed in Chapter 16, scientists have succeeded in sequencing virtually all the genetic information in a human cell, and the genetic information of many other organisms is being studied. Some people might think that the sequencing of genes is the end of the story, but it is actually only the beginning. Much genetic information is used to specify the structure of proteins, and biologists may devote most of the 21st century to understanding these extraordinarily multifaceted macromolecules that are of central importance in the chemistry of life. In a real sense, proteins are involved in virtually all aspects of metabolism because most enzymes, molecules that accelerate the thousands of different chemical reactions that take place in an organism, are proteins. Proteins are assembled into a variety of shapes, allowing them to serve as major structural components of cells and tissues. For this reason, growth and repair, as well as maintenance of the organism, depend on proteins. As shown in TABLE 3-2, proteins perform many other functions. Each cell type contains characteristic forms, distributions, and amounts of protein that largely determine what the cell looks like and how it functions. A muscle cell contains large amounts of the proteins myosin and actin, which are responsible for its appearance as well as its ability to contract. The protein hemoglobin, found in red blood cells, is responsible for the specialized function of oxygen transport.

7 8

Amino acids are the subunits of proteins

FIGURE 3-15 Steroids Four attached rings—three six-carbon rings and one with five carbons— make up the fundamental structure of a steroid (shown in green). Note that some carbons are shared by two rings. In these simplified structures, a carbon atom is present at each angle of a ring; the hydrogen atoms attached directly to the carbon atoms have not been drawn. Steroids are mainly distinguished by their attached functional groups.

the juvenile hormone of insects, are also fatty acid derivatives (discussed in Chapter 49).

Review ■

■

Why do saturated, unsaturated, and trans fatty acids differ in their properties? Why do phospholipids form lipid bilayers in aqueous conditions?

3.4 PROTEINS

9

Give an overall description of the structure and functions of proteins. Describe the features that are shared by all amino acids and explain how amino acids are grouped into classes based on the characteristics of their side chains. Distinguish among the four levels of organization of protein molecules.

Amino acids, the constituents of proteins, have an amino group (ONH2) and a carboxyl group (OCOOH) bonded to the same

H H

N H

C CH3

O C

H OH

H

H

N+

C

H

CH3

O

and histidine. Arginine is added to the list for children because they do not synthesize enough to support growth.

C O–

Ionized form

FIGURE 3-16 An amino acid at pH 7 In living cells, amino acids exist mainly in their ionized form, as dipolar ions.

asymmetrical carbon atom, known as the alpha carbon. Amino acids in solution at neutral pH are mainly dipolar ions; that is, they possess a positive charge at one end and a negative charge at the opposite end. This is generally how amino acids exist at cell pH. Each carboxyl group (OCOOH) donates a proton and becomes ionized (OCOO−), whereas each amino group (ONH2) accepts a proton and becomes ONH3+ (FIG. 3-16). Because of the ability of their amino and carboxyl groups to accept and release protons, amino acids in solution resist changes in acidity and alkalinity and therefore are important biological buffers. Twenty amino acids are commonly found in proteins, each identified by the variable side chain (R group) bonded to the a carbon (FIG. 3-17). Glycine, the simplest amino acid, has a hydrogen atom as its R group; alanine has a methyl (OCH3) group. The amino acids are grouped in Figure 3-17 by the properties of their side chains. These broad groupings actually include amino acids with a fairly wide range of properties. Amino acids classified as having nonpolar side chains tend to have hydrophobic properties, whereas those classified as polar are more hydrophilic. An acidic amino acid has a side chain that contains a carboxyl group. At cell pH the carboxyl group is dissociated, giving the R group a negative charge. A basic amino acid becomes positively charged when the amino group in its side chain accepts a hydrogen ion. Acidic and basic side chains are ionic at cell pH and therefore hydrophilic. Some proteins have unusual amino acids in addition to the 20 common ones. These rare amino acids are produced by the modification of common amino acids after they have become part of a protein. For example, after they have been incorporated into collagen, lysine and proline may be converted to hydroxylysine and hydroxyproline. These amino acids can form cross links between the peptide chains that make up collagen. Such cross links produce the firmness and great strength of the collagen molecule, which is a major component of cartilage, bone, and other connective tissues. With some exceptions, prokaryotes and plants synthesize all their needed amino acids from simpler substances. If the proper raw materials are available, the cells of animals can manufacture some, but not all, of the biologically significant amino acids. Essential amino acids are those an animal cannot synthesize in amounts sufficient to meet its needs and must obtain from the diet. Animals differ in their biosynthetic capacities; what is an essential amino acid for one species may not be for another. The essential amino acids for humans are isoleucine, leucine, lysine, methionine, phenylalanine, threonine, tryptophan, valine,

Peptide bonds join amino acids Amino acids combine chemically with one another by a condensation reaction that bonds the carboxyl carbon of one molecule to the amino nitrogen of another (FIG. 3-18). The covalent carbon-tonitrogen bond linking two amino acids is a peptide bond. When two amino acids combine, a dipeptide is formed; a longer chain of amino acids is a polypeptide. A protein consists of one or more polypeptide chains. Each polypeptide has a free amino group at one end and a free carboxyl group (belonging to the last amino acid added to the chain) at the opposite end. The other amino and carboxyl groups of the amino acid monomers (except those in side chains) are part of the peptide bonds. The process by which polypeptides are synthesized is discussed in Chapter 13. A polypeptide may contain hundreds of amino acids joined in a specific linear order. The backbone of the polypeptide chain includes the repeating sequence N

C

C

N

C

C

N

C

C

plus all other atoms except those in the R groups. The R groups of the amino acids extend from this backbone. An almost infinite variety of protein molecules is possible, differing from one another in the number, types, and sequences of amino acids they contain. The 20 types of amino acids found in proteins may be thought of as letters of a protein alphabet; each protein is a very long sentence made up of amino acid letters.

Proteins have four levels of organization The polypeptide chains making up a protein are twisted or folded to form a macromolecule with a specific conformation, or 3-D shape. Some polypeptide chains form long fibers. Globular proteins are tightly folded into compact, roughly spherical shapes. There is a close relationship between a protein’s conformation and its function. For example, a typical enzyme is a globular protein with a unique shape that allows it to catalyze a specific chemical reaction. Similarly, the shape of a protein hormone enables it to combine with receptors on its target cell (the cell on which the hormone acts). Scientists recognize four main levels of protein organization: primary, secondary, tertiary, and quaternary. Primary structure is the amino acid sequence The sequence of amino acids, joined by peptide bonds, is the primary structure of a polypeptide chain. As discussed in Chapter 13, this sequence is specified by the instructions in a gene. Using analytical methods, investigators can determine the exact sequence of amino acids in a protein molecule. The primary structures of thousands of proteins are known. For example, glucagon, a hormone secreted by the pancreas, is a small polypeptide, consisting of only 29 amino acid units (FIG. 3-19). Primary structure is always represented in a simple, linear, “beads-on-a-string” form. However, the overall conformation

(a) Nonpolar (hydrophobic) COO– H3N+

C

COO–

H

H2N+

CH2

H2C

CH3

Leucine (Leu, L)

Proline (Pro, P)

COO– H3N+

H

CH2 CH2

CH H3C

C

COO– H3N+

H

C

C

H

CH

CH3

CH3

Alanine (Ala, A)

CH3

Valine (Val, V)

(b) Polar, uncharged COO–

COO– H3N+

C

H

H

C

OH

H3N+

H

C CH2

CH3

OH Serine (Ser, S)

Threonine (Thr, T)

COO– COO– H3N+

C

H3N+

H

H

CH2

CH2

CH2

C O

C

C

NH2

O

Asparagine (Asn, N)

NH2

Glutamine (Gln, Q)

(c) Acidic COO– COO– H3N+

C

H3N+

H

COO–

Aspartic acid (Asp, D)

H

CH2

CH2

–

C

CH2

–

COO–

Glutamic acid (Glu, E)

COO– H3N+

C

COO–

H

H3N+

C

CH2

CH2

CH2 S

C CH

N H

CH3 Methionine (Met, M)

Tryptophan (Trp, W)

COO– H3N+

H

C

COO–

H

CH2

H3N+

C

H

H3C

C

H

CH2 CH3 Phenylalanine (Phe, F)

Isoleucine (Ile, I) COO–

COO– H3N+

C

H3N+

H

C

H

CH2

H

SH Glycine (Gly, G)

Cysteine (Cys, C) COO–

COO– H3N+

C

H3N+

H

H

CH2

CH2

HC H+N

OH

C

NH

C H

+

Tyrosine (Tyr, Y)

C

Histidine (His, H)

(d) Basic COO–

COO– H3N+

C

H3N+

H

CH2

CH2

CH2

CH2 CH2

CH2

NH

CH2 + Lysine (Lys, K)

H

C

NH3+

C +

FIGURE 3-17 The 20 common amino acids (a) Nonpolar amino acids have side chains that are relatively hydrophobic, whereas (b) polar amino acids have relatively hydrophilic side chains. Carboxyl groups and amino groups are electrically charged at cell pH; therefore, (c) acidic and (d) basic amino acids are hydrophilic. The standard three-letter and one-letter

H2

+N

NH2

Arginine (Arg, R)

R group H

H N H

Carboxyl group

C H

Amino group

CH3

H

O +

C

R group

OH

Glycine

N

C

H

Peptide bond O

H

C

H

N OH

Alanine

H

H

O

C

C

H

CH3 N H

C H

O C

+

H2O

OH

Glycylalanine (a dipeptide)

FIGURE 3-18 Animated Peptide bonds A dipeptide is formed by a condensation reaction, that is, by the removal of the equivalent of a water molecule from the carboxyl group of one amino acid and the amino group of another amino acid. The resulting peptide bond is a covalent, carbon-to-nitrogen bond. Note that the carbon is also part of a carbonyl group and that the nitrogen is also covalently bonded to a hydrogen. Additional amino acids can be added to form a long polypeptide chain with a free amino group at one end and a free carboxyl group at the other.

of a protein is far more complex, involving interactions among the various amino acids that make up the primary structure of the molecule. Therefore, the higher orders of structure—secondary, tertiary, and quaternary—ultimately derive from the specific amino acid sequence (the primary structure). Secondary structure results from hydrogen bonding involving the backbone Some regions of a polypeptide exhibit secondary structure, which is highly regular. The two most common types of secondary structure are the a-helix and the b-pleated sheet; the designations a and b refer simply to the order in which these two types of secondary structure were discovered. An a-helix is a region where a polypeptide chain forms a uniform helical coil (FIG. 3-20a). The helical structure is determined and maintained by the formation of hydrogen bonds between the backbones of the amino acids in successive turns of the spiral coil. Each hydrogen bond forms between an oxygen with a partial negative charge and a hydrogen with a partial positive charge. The oxygen is part of the remnant of the carboxyl group of one amino acid; the hydrogen is part of the remnant of the amino group of the fourth amino acid down the chain. Thus, 3.6 amino acids are included in each complete turn of the helix. Every amino acid in an a-helix is hydrogen bonded in this way. The a-helix is the basic structural unit of some fibrous proteins that make up wool, hair, skin, and nails. The elasticity of these fibers is due to a combination of physical factors (the helical shape) and chemical factors (hydrogen bonding). Although hydrogen bonds maintain the helical structure, these bonds can be broken, allowing the fibers to stretch under tension (like a telephone cord). When

+

the tension is released, the fibers recoil and hydrogen bonds reform. This explains why you can stretch the hairs on your head to some extent and they will snap back to their original length. The hydrogen bonding in a b-pleated sheet takes place between different polypeptide chains, or different regions of a polypeptide chain that has turned back on itself (FIG. 3-20b). Each chain is fully extended; however, because each has a zigzag structure, the resulting “sheet” has an overall pleated conformation (much like a sheet of paper that has been folded to make a fan). Although the pleated sheet is strong and flexible, it is not elastic, because the distance between the pleats is fixed, determined by the strong covalent bonds of the polypeptide backbones. Fibroin, the protein of silk, is characterized by a b-pleated sheet structure, as are the cores of many globular proteins. It is not uncommon for a single polypeptide chain to include both a-helical regions and regions with b-pleated sheet conformations. The properties of some biological materials result from such combinations. A spider’s web is composed of a material that is extremely strong, flexible, and elastic. Once again we see function and structure working together, as these properties derive from the fact that spider silk is a composite of proteins with a-helical conformations (providing elasticity) and others with b-pleated sheet conformations (providing strength). Tertiary structure depends on interactions among side chains The tertiary structure of a protein molecule is the overall shape assumed by each individual polypeptide chain (FIG. 3-21). This 3-D structure is determined by four main factors that involve interactions among R groups (side chains) belonging to the same

H3N His Ser Gln Gly Thr Phe Thr Ser Asp Tyr Ser Lys Tyr Leu Asp Ser Arg Arg Ala Gln Asp Phe Val Gln Trp Leu Met Asn Thr COO– 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29

FIGURE 3-19 Primary structure of a polypeptide Glucagon is a very small polypeptide made up of 29 amino acids. The linear sequence of amino acids is indicated by ovals containing their abbreviated names (see Fig. 3-17).

KEY POINT

Secondary structure is highly regular.

KEY: Carbon atom

C

Oxygen atom

C

Nitrogen atom

N C

Hydrogen atom

C N

H C

C

N O

Hydrogen bonds hold helix coils in shape C

R group

C

N

N

C

C

C

(a) In an Ĝ-helix the R groups project out from the sides. (The R groups have been omitted in the simplified diagram at left.)

Hydrogen bonds hold neighboring strands of sheet together

(b) A ĝ-pleated sheet forms when a polypeptide chain folds back on itself (arrows); half the R groups project above the sheet, and the other half project below it.

FIGURE 3-20 Animated Secondary structure of a protein

polypeptide chain. These include both weak interactions (hydrogen bonds, ionic bonds, and hydrophobic interactions) and strong covalent bonds.

hydrogens are removed, and the two sulfur atoms that remain become covalently linked.

1. Hydrogen bonds form between R groups of certain amino acid subunits. 2. An ionic bond can occur between an R group with a unit of positive charge and one with a unit of negative charge. 3. Hydrophobic interactions result from the tendency of nonpolar R groups to be excluded by the surrounding water and therefore to associate in the interior of the globular structure. 4. Covalent bonds known as disulfide bonds or disulfide bridges (OSOSO) may link the sulfur atoms of two cysteine subunits belonging to the same chain. A disulfide bridge forms when the sulfhydryl groups of two cysteines react; the two

Quaternary structure results from interactions among polypeptides Many functional proteins are composed of two or more polypeptide chains, interacting in specific ways to form the biologically active molecule. Quaternary structure is the resulting 3-D structure. The same types of interactions that produce secondary and tertiary structure also contribute to quaternary structure; these include hydrogen bonding, ionic bonding, hydrophobic interactions, and disulfide bridges. A functional antibody molecule, for example, consists of four polypeptide chains joined by disulfide bridges (see Chapter 45).

KEY POINT

Tertiary structure depends on side chain interactions.

Ionic bond

Hydrogen bond

CH2 C O CH2

HO

NH+3 –O O

C CH2

CH2

H3C

O H

S

CH

O

S

H3C

C

CH3 HC CH3

CH2

CH2

Hydrophobic interaction Jane Richardson

Disulfide bond

(a) Hydrogen bonds, ionic bonds, hydrophobic interactions, and disulfide bridges between R groups hold the parts of the molecule in the designated shape.

(b) In this drawing, α-helical regions are represented by purple coils, β-pleated sheets by broad green ribbons, and connecting regions by narrow tan ribbons. The interactions among R groups that stabilize the bends and foldbacks that give the molecule its overall conformation (tertiary structure) are represented in yellow. This protein is bovine ribonuclease a.

FIGURE 3-21 Animated Tertiary structure of a protein

Disulfide bridges are a common feature of antibodies and other proteins secreted from cells. These strong bonds stabilize the molecules in the extracellular environment. Hemoglobin, the protein in red blood cells responsible for oxygen transport, is an example of a globular protein with a quaternary structure (FIG. 3-22a). Hemoglobin consists of 574 amino acids arranged in four polypeptide chains: two identical chains called alpha chains and two identical chains called beta chains. Collagen, mentioned previously, has a fibrous type of quaternary structure that allows it to function as the major strengthener of animal tissues. It consists of three polypeptide chains wound about one another and bound by cross links between their amino acids (FIG. 3-22b).

The amino acid sequence of a protein determines its conformation In 1972, U.S. researcher Christian B. Anfinsen was awarded the Nobel Prize in Chemistry for his studies on protein folding, which

demonstrated that, at least under defined experimental conditions in vitro (outside a living cell), a polypeptide can spontaneously undergo folding processes that yield its normal, functional conformation. Since Anfinsen’s pioneering work, many researchers studying various proteins and using a variety of highly sophisticated approaches have amassed evidence supporting the widely held conclusion that amino acid sequence is the ultimate determinant of protein conformation. However, because conditions in vivo (in the cell) are quite different from defined laboratory conditions, proteins do not always fold spontaneously in the cell. On the contrary, scientists have learned that proteins known as molecular chaperones mediate the folding of other protein molecules. Molecular chaperones are thought to make the folding process more orderly and efficient and to prevent partially folded proteins from becoming inappropriately aggregated. However, there is no evidence that molecular chaperones actually dictate the folding pattern. For this reason, the existence of chaperones is not an argument against the idea that amino acid sequence determines conformation.

KEY POINT

Heme

ing a more random conformation. This unfolding, which is mainly due to the disruption of hydrogen bonds and ionic bonds, is typically accompanied by a loss of normal function. Such changes in shape and the accompanying loss of biological activity are termed denaturation of the protein. For example, a denatured enzyme would lose its ability to catalyze a chemical reaction. An everyday example of denaturation occurs when we fry an egg. The consistency of the egg white protein, known as albumin, changes to a solid. Denaturation generally cannot be reversed (you cannot “unfry” an egg). However, under certain conditions, some proteins have been denatured and have returned to their original shape and biological activity when normal environmental conditions were restored.

Proteins with two or more polypeptide chains have quaternary structure. Beta chain (β-globin)

Alpha chain (α-globin)

Alpha chain (α-globin)

Beta chain (β-globin)

(a) Hemoglobin, a globular protein, consists of four polypeptide chains, each joined to an iron-containing molecule, a heme.

(b) Collagen, a fibrous protein, is a triple helix consisting of three long polypeptide chains.

FIGURE 3-22 Quaternary structure of a protein

Protein conformation determines function The overall structure of a protein helps determine its biological activity. A single protein may have more than one distinct structural region, called a domain, each with its own function. Many proteins are modular, consisting of two or more globular domains, connected by less compact regions of the polypeptide chain. Each domain may have a different function. For example, a protein may have one domain that attaches to a membrane and another that acts as an enzyme. The biological activity of a protein can be disrupted by a change in amino acid sequence that results in a change in conformation. For example, the genetic disease known as sickle cell anemia is due to a mutation that causes the substitution of the amino acid valine for glutamic acid at position 6 (the sixth amino acid from the amino end) in the beta chain of hemoglobin. The substitution of valine (which has a nonpolar side chain) for glutamic acid (which has a charged side chain) makes the hemoglobin less soluble and more likely to form crystal-like structures. This alteration of the hemoglobin affects the red blood cells, changing them to the crescent or sickle shapes that characterize this disease (see Fig. 16-8). The biological activity of a protein may be affected by changes in its 3-D structure. When a protein is heated, subjected to significant pH changes, or treated with certain chemicals, its structure becomes disordered and the coiled peptide chains unfold, yield-

Protein conformation is studied through a variety of methods The architecture of a protein can be ascertained directly through various types of analysis, such as the X-ray diffraction studies discussed in Chapter 12. Because these studies are tedious and costly, researchers are developing alternative approaches, which rely heavily on the enormous databases generated by the Human Genome Project and related initiatives. Today a protein’s primary structure can be determined rapidly through the application of genetic engineering techniques (discussed in Chapter 15) or by the use of sophisticated technology such as mass spectrometry. Researchers use these amino acid sequence data to predict a protein’s higher levels of structure. As you have seen, side chains interact in relatively predictable ways, such as through ionic and hydrogen bonds. In addition, regions with certain types of side chains are more likely to form a-helices or b-pleated sheets. Computer programs make such predictions, but these are often imprecise because of the many possible combinations of folding patterns. Computers are an essential part of yet another strategy. Once the amino acid sequence of a polypeptide has been determined, researchers use computers to search databases to find polypeptides with similar sequences. If the conformations of any of those polypeptides or portions have already been determined directly by X-ray diffraction or other techniques, this information can be extrapolated to make similar correlations between amino acid sequence and 3-D structure for the protein under investigation. These predictions are increasingly reliable, as more information is added to the databases every day.

Misfolded proteins are implicated in human diseases Studies on the mechanisms of protein folding, and on the relationship between the activity of a protein and its conformation, are increasingly of medical importance. For example, as discussed in Chapter 24, mad cow disease and related diseases in humans and other animals are caused by misfolded proteins called prions. Other serious diseases in which misfolded proteins play an important role include Alzheimer’s disease (see Chapter 41) and Huntington’s disease (see Chapter 16).

Review ■

■

Draw the structural formula of a simple amino acid. What is the importance of the carboxyl group, amino group, and R group? How does the primary structure of a polypeptide influence its secondary and tertiary structures?

O

NH2 N O

CH

HN

CH N H Cytosine (C)

O

O CH3

C

CH N H Thymine (T)

O

HN

CH

CH N H Uracil (U)

(a) Pyrimidines. The three major pyrimidine bases found in nucleotides are cytosine, thymine (in DNA only), and uracil (in RNA only).

NH2

O N

N

N

HN CH

HC

3.5 NUCLEIC ACIDS ■ ■ LEARNING OBJECTIVE 10 Describe the components of a nucleotide. Name some nucleic acids and nucleotides, and discuss the importance of these compounds in living organisms.

N H Adenine (A) N

CH H2N

N

N H

Guanine (G)

(b) Purines. The two major purine bases found in nucleotides are adenine and guanine.

FIGURE 3-23 Animated Components of nucleotides

Nucleic acids transmit hereditary information and determine what proteins a cell manufactures. Two classes of nucleic acids are found in cells: deoxyribonucleic acid and ribonucleic acid. Deoxyribonucleic acid (DNA) composes the genes, the hereditary material of the cell, and contains instructions for making all the proteins, as well as all the RNA the organism needs. Ribonucleic acid (RNA) participates in the process in which amino acids are linked to form polypeptides. Some types of RNA, known as ribozymes, can even act as specific biological catalysts. Like proteins, nucleic acids are large, complex molecules. The name nucleic acid reflects the fact that they are acidic and were first identified, by Swiss biochemist Friedrich Miescher in 1870, in the nuclei of pus cells. Nucleic acids are polymers of nucleotides, molecular units that consist of (1) a five-carbon sugar, either deoxyribose (in DNA) or ribose (in RNA); (2) one or more phosphate groups, which make the molecule acidic; and (3) a nitrogenous base, a ring compound that contains nitrogen. The nitrogenous base may be either a double-ring purine or a single-ring pyrimidine (FIG. 3-23). DNA commonly contains the purines adenine (A) and guanine (G), the pyrimidines cytosine (C) and thymine (T), the sugar deoxyribose, and phosphate. RNA contains the purines adenine and guanine, and the pyrimidines cytosine and uracil (U), together with the sugar ribose, and phosphate. The molecules of nucleic acids are made of linear chains of nucleotides, which are joined by phosphodiester linkages, each consisting of a phosphate group and the covalent bonds that attach it to the sugars of adjacent nucleotides (FIG. 3-24). Note that each nucleotide is defined by its particular base and that nucleotides can be joined in any sequence. A nucleic acid molecule is uniquely defined by its specific sequence of nucleotides, which constitutes a kind of code (see

The hydrogens indicated by the white boxes are removed when the base is attached to a sugar.

Chapter 13). Whereas RNA is usually composed of one nucleotide chain, DNA consists of two nucleotide chains held together by hydrogen bonds and entwined around each other in a double helix (see Fig. 1-7).

Some nucleotides are important in energy transfers and other cell functions In addition to their importance as subunits of DNA and RNA, nucleotides perform other vital functions in living cells. Adenosine triphosphate (ATP), composed of adenine, ribose, and three phosphates (see Fig. 7-5), is of major importance as the primary energy currency of all cells (see Chapter 7). The two terminal phosphate groups are joined to the nucleotide by covalent bonds. These are traditionally indicated by wavy lines, which indicate that ATP can transfer a phosphate group to another molecule, making that molecule more reactive. In this way ATP is able to donate some of its chemical energy. Most of the readily available chemical energy of the cell is associated with the phosphate groups of ATP. Like ATP, guanosine triphosphate (GTP), a nucleotide that contains the base guanine, can transfer energy by transferring a phosphate group and also has a role in cell signaling (see Chapter 6). A nucleotide may be converted to an alternative form with specific cell functions. ATP, for example, is converted to cyclic adenosine monophosphate (cyclic AMP, or cAMP) by the enzyme adenylyl cyclase (FIG. 3-25). Cyclic AMP regulates certain

NH2 C

O

5′ –O

O

P O

Nucleotide

CH2

N

O

O –O

O

Uracil

O

5′ CH2

H

OH O

P

–

O

NH2

O

N

CH2

N

O

O

H O

P

N

H 3′

H

OH

O

N

Cyclic AMP

N Adenine

Ribose

C

N

N

Ribose

CH

HC

O

N

C

N

FIGURE 3-25 Cyclic adenosine monophosphate (cAMP) The single phosphate is part of a ring connecting two regions of the ribose.

Phosphodiester linkage

O –O

OH

NH2

O

P

N

O CH2 O

O Cytosine

N

Ribose O –O

ter 7, nicotinamide adenine dinucleotide has a primary role in oxidation and reduction reactions in cells. It can exist in an oxidized form (NAD+) that is converted to a reduced form (NADH) when it accepts electrons (in association with hydrogen; see Fig. 7-7). These electrons, along with their energy, are transferred to other molecules.

O

OH O

N

O CH2 O

N

P

Ribose OH 3′

Review N

N Guanine

■

How does the structure of a nucleotide relate to its function?

NH2

OH

3.6 IDENTIFYING BIOLOGICAL MOLECULES ■ ■ LEARNING OBJECTIVE 11 Compare the functions and chemical compositions of the major groups of organic compounds: carbohydrates, lipids, proteins, and nucleic acids.

FIGURE 3-24 RNA, a nucleic acid Nucleotides, each with a specific base, are joined by phosphodiester linkages.

cell functions, such as cell signaling, and is important in the mechanism by which some hormones act. A related molecule, cyclic guanosine monophosphate (cGMP), also plays a role in certain cell signaling processes. Cells contain several dinucleotides, which are of great importance in metabolic processes. For example, as discussed in Chap-

Although the classes of biological molecules may seem overwhelming at first, you will learn to distinguish them readily by understanding their chief attributes. These are summarized in TABLE 3-3.

Review ■

How can you distinguish a pentose sugar from a hexose sugar? a disaccharide from a sterol? an amino acid from a monosaccharide? a phospholipid from a triacylglycerol? a protein from a polysaccharide? a nucleic acid from a protein?

TABLE 3-3 Class and Component Elements Carbohydrates C, H, O

Lipids C, H, O (sometimes N, P)

Classes of Biologically Important Organic Compounds

Description

How to Recognize

Principal Function in Living Systems

Contain approximately 1 C:2 H:1 O (but make allowance for loss of oxygen when sugar units as nucleic acids and glycoproteins are linked)

Count the carbons, hydrogens, and oxygens.

Cell fuel; energy storage; structural component of plant cell walls; component of other compounds such as nucleic acids and glycoproteins

1. Monosaccharides (simple sugars). Mainly fivecarbon (pentose) molecules such as ribose or six-carbon (hexose) molecules such as glucose and fructose

Look for the ring shapes:

Cell fuel; components of other compounds

2. Disaccharides. Two sugar units linked by a glycosidic bond, e.g., maltose, sucrose

Count sugar units.

Components of other compounds; form of sugar transported in plants

3. Polysaccharides. Many sugar units linked by glycosidic bonds, e.g., glycogen, cellulose

Count sugar units.

Energy storage; structural components of plant cell walls

Contain much less oxygen relative to carbon and hydrogen than do carbohydrates 1. Fats. Combination of glycerol with one to three fatty acids. Monoacylglycerol contains one fatty acid; diacylglycerol contains two fatty acids; triacylglycerol contains three fatty acids. If fatty acids contain double carbon-to-carbon linkages (CAC), they are unsaturated; otherwise, they are saturated.

Energy storage; cellular fuel, components of cells; thermal insulation Look for glycerol at one end of molecule: H H

C

O

H

C

O

H

C

O

Cell fuel; energy storage

H 2. Phospholipids. Composed of glycerol attached to one or two fatty acids and to an organic base containing phosphorus

Look for glycerol and side chain containing phosphorus and nitrogen.

Components of cell membranes

3. Steroids. Complex molecules containing carbon atoms arranged in four attached rings (Three rings contain six carbon atoms each, and the fourth ring contains five.)

Look for four attached rings:

Some are hormones; others include cholesterol, bile salts, vitamin D; components of cell membranes

4. Carotenoids. Orange and yellow pigments; consist of isoprene units

Look for isoprene units. H CH3

Converted to retinal (important in photoreception) and vitamin A

H2C

C

C

CH2

Proteins C, H, O, N (usually S)

One or more polypeptides (chains of amino acids) coiled or folded in characteristic shapes

Look for amino acid units joined by CON bonds.

Serve as enzymes; structural components; muscle proteins; hemoglobin

Nucleic Acids C, H, O, N, P

Backbone composed of alternating pentose and phosphate groups, from which nitrogenous bases project. DNA contains the sugar deoxyribose and the bases guanine, cytosine, adenine, and thymine. RNA contains the sugar ribose and the bases guanine, cytosine, adenine, and uracil. Each molecular subunit, called a nucleotide, consists of a pentose, a phosphate, and a nitrogenous base.

Look for a pentose-phosphate backbone. DNA forms a double helix.

Storage, transmission, and expression of genetic information; some important in energy transfers, cell signaling, and other aspects of metabolism.

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

3.1 (page 47)

glycogen. The cell walls of plants are composed mainly of the structural polysaccharide cellulose.

1 Describe the properties of carbon that make it the central component of organic compounds. Each carbon atom forms four covalent bonds with up to four other atoms; these bonds are single, double, or triple bonds. Carbon atoms form straight or branched chains or join into rings. Carbon forms covalent bonds with a greater number of different elements than does any other type of atom. 2 Define the term isomer and distinguish among the three principal isomer types. ■ Isomers are compounds with the same molecular formula but different structures. ■ Structural isomers differ in the covalent arrangements of their atoms. Geometric isomers, or cis–trans isomers, differ in the spatial arrangements of their atoms. Enantiomers are isomers that are mirror images of each other. Cells can distinguish between these configurations. 3 Identify the major functional groups present in organic compounds and describe their properties. Hydrocarbons, organic compounds consisting of only carbon and ■ hydrogen, are nonpolar and hydrophobic. The methyl group is a hydrocarbon group. ■ Polar and ionic functional groups interact with one another and are hydrophilic. Partial charges on atoms at opposite ends of a bond are responsible for the polar property of a functional group. Hydroxyl and carbonyl groups are polar. ■ Carboxyl and phosphate groups are acidic, becoming negatively charged when they release hydrogen ions. The amino group is basic, becoming positively charged when it accepts a hydrogen ion. 4 Explain the relationship between polymers and macromolecules. ■ Long chains of monomers (similar organic compounds) linked through condensation reactions are called polymers. Large polymers such as polysaccharides, proteins, and DNA are referred to as macromolecules. They can be broken down by hydrolysis reactions.

Learn more about starch and cellulose by clicking on the figure in CengageNOW.

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3.3 (page 56) 6 Distinguish among fats, phospholipids, and steroids, and describe the composition, characteristics, and biological functions of each. ■ Lipids are composed mainly of hydrocarbon-containing regions, with few oxygen-containing (polar or ionic) functional groups. Lipids have a greasy or oily consistency and are relatively insoluble in water. ■ Triacylglycerol, the main storage form of fat in organisms, consists of a molecule of glycerol combined with three fatty acids. Monoacylglycerols and diacylglycerols contain one and two fatty acids, respectively. A fatty acid can be either saturated with hydrogen or unsaturated.

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Learn more about triacylglycerol and other lipids by clicking on the figures in CengageNOW.

3.4 (page 60) 7 Give an overall description of the structure and functions of proteins. ■

Learn more about condensation and hydrolysis reactions by clicking on the figure in CengageNOW.

3.2 (page 51) 5 Distinguish among monosaccharides, disaccharides, and polysaccharides. Compare storage polysaccharides with structural polysaccharides. ■ Carbohydrates contain carbon, hydrogen, and oxygen in a ratio of approximately one carbon to two hydrogens to one oxygen. Monosaccharides are simple sugars such as glucose, fructose, and ribose. Two monosaccharides join by a glycosidic linkage to form a disaccharide such as maltose or sucrose. CH2OH H

O H OH

HOCH2

H

HO

O H

OH

■

Learn more about amino acids and peptide bonds by clicking on the figures in CengageNOW.

8 Describe the features that are shared by all amino acids and explain how amino acids are grouped into classes based on the characteristics of their side chains. COO– H3N+

HO CH2OH

OH

Proteins are complex macromolecules made of simpler subunits, called amino acids, joined by peptide bonds. Two amino acids combine to form a dipeptide. A longer chain of amino acids is a polypeptide. Proteins are the most versatile class of biological molecules, serving a variety of functions, such as enzymes, structural components, and cell regulators. Proteins are composed of various linear sequences of 20 different amino acids.

H

O H

■

H

Phospholipids are structural components of cell membranes. A phospholipid consists of a glycerol molecule attached at one end to two fatty acids and at the other end to a phosphate group linked to an organic compound such as choline. Steroid molecules contain carbon atoms arranged in four attached rings. Cholesterol, bile salts, and certain hormones are important steroids.

H

Most carbohydrates are polysaccharides, long chains of repeating units of a simple sugar. Carbohydrates are typically stored in plants as the polysaccharide starch and in animals as the polysaccharide

C

H

CH3 ■

All amino acids contain an amino group and a carboxyl group. Amino acids vary in their side chains, which dictate their chemical properties—nonpolar, polar, acidic, or basic. Amino acids generally exist as dipolar ions at cell pH and serve as important biological buffers.

9 Distinguish among the four levels of organization of protein molecules. ■

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Primary structure is the linear sequence of amino acids in the polypeptide chain. Secondary structure is a regular conformation, such as an a-helix or a b-pleated sheet; it is due to hydrogen bonding between elements of the backbones of the amino acids. Tertiary structure is the overall shape of the polypeptide chains, as dictated by chemical properties and interactions of the side chains of specific amino acids. Hydrogen bonds, ionic bonds, hydrophobic interactions, and disulfide bridges contribute to tertiary structure. Quaternary structure is determined by the association of two or more polypeptide chains. Learn more about the structure of a protein by clicking on the figure in CengageNOW.

3.5 (page 68)

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The nucleic acids DNA and RNA, composed of long chains of nucleotide subunits, store and transfer information that specifies the sequence of amino acids in proteins and ultimately the structure and function of the organism. Nucleotides are composed of a two-ring purine or one-ring pyrimidine nitrogenous base, a five-carbon sugar (ribose or deoxyribose), and one or more phosphate groups. ATP (adenosine triphosphate) is a nucleotide of special significance in energy metabolism. NAD+ is also involved in energy metabolism through its role as an electron (hydrogen) acceptor in biological oxidation and reduction reactions.

3.6 (page 69) 11 Compare the functions and chemical compositions of the major groups of organic compounds: carbohydrates, lipids, proteins, and nucleic acids. ■ Review Table 3-3.

10 Describe the components of a nucleotide. Name some nucleic acids and nucleotides, and discuss the importance of these compounds in living organisms.

T E S T YO U R U N D E R S TA N D I N G 1. Carbon is particularly well suited to be the backbone of organic molecules because (a) it can form both covalent bonds and ionic bonds (b) its covalent bonds are very irregularly arranged in three-dimensional space (c) its covalent bonds are the strongest chemical bonds known (d) it can bond to atoms of a large number of other elements (e) all the bonds it forms are polar 2. The structures depicted are (a) enantiomers (b) different views of the same molecule (c) geometric (cis–trans) isomers (d) both geometric isomers and enantiomers (e) structural isomers CH3 CH3 H

C

C

H

H

H

H

H

CH3

C

C

H

CH3 H

3. Which of the following is a nonpolar molecule? (a) water, H2O (b) ammonia, NH3 (c) methane, CH4 (d) ethane, C2H6 (e) more than one of the preceding 4. The synthetic process by which monomers are covalently linked is (a) hydrolysis (b) isomerization (c) condensation (d) glycosidic linkage (e) ester linkage 5. A monosaccharide designated as an aldehyde sugar contains (a) a terminal carboxyl group (b) an internal carboxyl group (c) a terminal carbonyl group (d) an internal carbonyl group (e) a terminal carboxyl group and an internal carbonyl group 6. Structural polysaccharides typically (a) have extensive hydrogen bonding between adjacent molecules (b) are much more hydrophilic than storage polysaccharides (c) have much stronger covalent bonds than do storage polysaccharides (d) consist of alternating a-glucose and b-glucose subunits (e) form helical structures in the cell

7. Saturated fatty acids are so named because they are saturated with (a) hydrogen (b) water (c) hydroxyl groups (d) glycerol (e) double bonds 8. Fatty acids in phospholipids and triacylglycerols interact with one another by (a) disulfide bridges (b) van der Waals interactions (c) covalent bonds (d) hydrogen bonds (e) fatty acids do not interact with one another 9. Which pair of amino acid side groups would be most likely to associate with each other by an ionic bond? 1. OCH3 2. OCH2OCOO− 3. OCH2OCH2ONH3+ 4. OCH2OCH2OCOO− 5. OCH2OOH (a) 1 and 2 (b) 2 and 4 (c) 1 and 5 (d) 2 and 5 (e) 3 and 4 10. Which of the following levels of protein structure may be affected by hydrogen bonding? (a) primary and secondary (b) primary and tertiary (c) secondary, tertiary, and quaternary (d) primary, secondary, and tertiary (e) primary, secondary, tertiary, and quaternary 11. Which of the following associations between R groups are the strongest? (a) hydrophobic interactions (b) hydrogen bonds (c) ionic bonds (d) peptide bonds (e) disulfide bridges 12. Each phosphodiester linkage in DNA or RNA includes a phosphate joined by covalent bonds to (a) two bases (b) two sugars (c) two additional phosphates (d) a sugar, a base, and a phosphate (e) a sugar and a base

CRITICAL THINKING 1. Like oxygen, sulfur forms two covalent bonds. However, sulfur is far less electronegative. In fact, it is approximately as electronegative as carbon. How would the properties of the various classes of biological molecules be altered if you were to replace all the oxygen atoms with sulfur atoms? 2. Hydrogen bonds and van der Waals interactions are much weaker than covalent bonds, yet they are vital to organisms. Why? 3. EVOLUTION LINK. In what ways are all species alike biochemically? Why? Identify some ways in which species may differ from one another biochemically. Why? 4. EVOLUTION LINK. The total number of possible amino acid sequences in a polypeptide chain is staggering. Given that there are 20 amino acids, potentially there could be 20100 different

amino acid sequences (an impossibly large number), just for polypeptides only 100 amino acids in length. However, the actual number of different polypeptides occurring in organisms is only a tiny fraction of this. Why? 5. EVOLUTION LINK. Each amino acid could potentially exist as one of two possible enantiomers, known as the d-form and the l-form (based on the arrangement of the groups attached to the asymmetric a carbon). However, in all organisms, only l-amino acids are found in proteins. What does this suggest about the evolution of proteins? Additional questions are available in CengageNOW at www.cengage.com/ login.

Organization of the Cell

Jennifer C. Waters/Photo Researchers, Inc.

4

KEY CONCEPTS

The cytoskeleton. The cell shown here was stained with fluorescent antibodies (specific proteins) that bind to proteins associated with DNA (purple) and to a protein (tubulin) in microtubules (green). Microfilaments (red) are also visible. This type of microscopy, known as confocal fluorescence microscopy, shows the extensive distribution of microtubules in this cell.

T

he cell is the smallest unit that can carry out all activities we associate with life. When provided with essential nutrients and an appropriate

4.1 The cell is the basic unit of life; its organization and size are critical in maintaining homeostasis, and its size and shape are adapted for its function.

environment, some cells can be kept alive and growing in the laboratory

4.2 Biologists study cells using microscopes and biochem-

biology in Chapter 1. Even as we describe individual components of cells,

ical techniques such as cell fractionation.

4.3 Unlike prokaryotic cells, eukaryotic cells have internal membranes that divide the cell into compartments, allowing cells to conduct specialized activities within separate, small areas.

for many years. By contrast, no isolated part of a cell is capable of sustained survival. As you read this chapter, recall the discussion of systems we discuss how these components work together, generating complex biological systems within the cell. The cell itself is a highly intricate biological system, and groups of cells make up tissues, organs, and organisms. Each of these is a biological system.

4.4 In eukaryotic cells, genetic information coded in DNA

Most prokaryotes and many protists and fungi consist of a single

is located in the nucleus, which is typically the most prominent organelle in the cell.

cell. In contrast, most plants and animals are composed of millions of

4.5 Among the many organelles in the cytoplasm are ribosomes, which synthesize proteins; endoplasmic reticulum and Golgi complexes, which process proteins; and mitochondria and chloroplasts, which convert energy from one form to another.

cells. Cells are the building blocks of complex multicellular organisms. Although they are basically similar, cells are also extraordinarily diverse and versatile. They are modified in a variety of ways to carry out specialized functions. The cell is composed of a vast array of inorganic and organic ions

4.6 The cytoskeleton is a dynamic internal framework

and molecules, including water, salts, carbohydrates, lipids, proteins,

that functions in various types of cell movement.

and nucleic acids. These molecules are organized to form the structures

4.7 Most eukaryotic cells are surrounded by a cell coat;

within the cell and its biochemical pathways. Genetic information is

in addition, many animal cells are surrounded by an extracellular matrix; cells of most bacteria, archaea, fungi, and plants are surrounded by a cell wall.

stored in DNA molecules and is faithfully replicated. This information is

passed to each new generation of cells during cell division. Information in DNA codes for specific proteins that, in turn, determine cell structure and function. Cells exchange materials and energy with the environment. All living cells need one or more sources of energy, but a cell rarely obtains energy in a form that is immediately usable. Cells convert energy from one form to another, and that energy is used to carry out various activities, ranging from mechanical work to chemical synthesis. Cells convert energy to a convenient form, usually

out that the ancestry of all the cells alive today can be traced back to ancient times. Evidence that all living cells have a common origin is provided by the basic similarities in their structures and in the molecules of which they are made. When we examine a variety of diverse organisms, ranging from simple bacteria to the most complex plants and animals, we find striking similarities at the cellular level. Careful studies of shared cell characteristics help us trace the evolutionary history of various organisms and furnish powerful evidence that all organisms alive today had a common origin.

chemical energy stored in adenosine triphosphate, or ATP (see Chapter 3). Both the chemical reactions that convert energy from one form to another and the mechanisms of information transfer are essentially the same in all cells, from those in bacteria to those of large, multicellular plants and animals. Such similarities suggest evolutionary relationships. Thanks to advances in technology, cell biologists use increasingly sophisticated tools in their search to better understand the structure and function of cells. For example, investigation of the cytoskeleton (cell skeleton), currently an active and exciting area of research, has been greatly enhanced by advances in microscopy. In the photomicrograph, we see the extensive distribution of microtubules in cells. Microtubules are key components of the cytoskeleton. They help maintain cell shape, function in cell movement, and facilitate transport of materials within the cell.

4.1 THE CELL: BASIC UNIT OF LIFE ■ ■ LEARNING OBJECTIVES 1 2 3

Describe the cell theory and relate it to the evolution of life. Summarize the relationship between cell organization and homeostasis. Explain the relationship between cell size and homeostasis.

Cells, the building blocks of organisms, are dramatic examples of the underlying unity of all living things.

The cell theory is a unifying concept in biology Two German scientists, botanist Matthias Schleiden in 1838 and zoologist Theodor Schwann in 1839, used inductive reasoning to conclude that all plants and animals consist of cells. These investigators used their own observations and those of other scientists to reach their conclusions. Later, Rudolf Virchow, another German scientist, observed cells dividing and giving rise to daughter cells. In 1855, Virchow proposed that new cells form only by the division of previously existing cells. The work of Schleiden, Schwann, and Virchow contributed greatly to the development of the cell theory, the unifying concept that (1) cells are the basic living units of organization and function in all organisms and (2) that all cells come from other cells. About 1880, another German biologist, August Weismann, added an important corollary to Virchow’s concept by pointing

The organization of all cells is basically similar The organization of cells and their small size allow them to maintain homeostasis, an appropriate internal environment. Cells experience constant changes in their environments, such as deviations in salt concentration, pH, and temperature. They must work continuously to restore and maintain the internal conditions that enable their biochemical mechanisms to function. In order for the cell to maintain homeostasis, its contents must be separated from the external environment. The plasma membrane is a structurally distinctive surface membrane that surrounds all cells. By making the interior of the cell an enclosed compartment, the plasma membrane allows the chemical composition of the cell to be different from that outside the cell. The plasma membrane serves as a selective barrier between the cell contents and the outer environment. Cells exchange materials with the environment and can accumulate needed substances and energy stores. Most cells have internal structures, called organelles, that are specialized to carry out metabolic activities, such as converting energy to usable forms, synthesizing needed compounds, and manufacturing structures necessary for functioning and reproduction. Each cell has genetic instructions coded in its DNA, which is concentrated in a limited region of the cell.

Cell size is limited Although their sizes vary over a wide range (FIG. 4-1), most cells are microscopic and must be measured by very small units. The basic unit of linear measurement in the metric system (see inside back cover) is the meter (m), which is just a little longer than a yard. A millimeter (mm) is 1/1000 of a meter and is about as long as the bar enclosed in parentheses (-). The micrometer (μm) is the most convenient unit for measuring cells. A bar 1 μm long is 1/1,000,000 (one millionth) of a meter, or 1/1000 of a millimeter—far too short to be seen with the unaided eye. Most of us have difficulty thinking about units that are too small to see, but it is helpful to remember that a micrometer has the same relationship to a millimeter that a millimeter has to a meter (1/1000). As small as it is, the micrometer is actually too large to measure most cell components. For this purpose biologists use the nanometer (nm), which is 1/1,000,000,000 (one billionth) of a meter, or 1/1000 of a micrometer. To mentally move down to the world of the nanometer, recall that a millimeter is 1/1000 of a meter, a micrometer is 1/1000 of a millimeter, and a nanometer is 1/1000 of a micrometer.

Mitochondrion

Red blood cells Chloroplast

0.1 nm

Chicken egg

Virus

Protein

Atom

Human egg

Typical bacteria

Nucleus

Amino acids Ribosomes

1 nm

10 nm

Smallest bacteria

100 nm

Epithelial cell

1 μm

10 μm

100 μm

Frog egg Some nerve cells

1 mm

10 mm

100 mm

Adult human

1m

10 m

Electron microscope Light microscope Human eye Measurements 1 meter 1 millimeter 1 micrometer

= = =

1000 millimeters (mm) 1000 micrometers (μm) 1000 nanometers (nm)

FIGURE 4-1 Biological size and cell diversity We can compare relative size from the chemical level to the level of an entire organism by using a logarithmic scale (multiples of 10). The prokaryotic cells of most bacteria range in size from 1 to 10 µm long. Most eukaryotic cells are between 10 and 30 µm in diameter. Mitochondria are about the size of small bacteria, whereas chloroplasts are usually larger, about 5 µm long. Ova (egg cells) are among the largest cells. Although microscopic, some nerve cells are very long. The cells shown here are not drawn to scale.

A few specialized algae and animal cells are large enough to be seen with the naked eye. A human egg cell, for example, is about 130 μm in diameter, or approximately the size of the period at the end of this sentence. The largest cells are birds’ eggs, but they are not typical cells because they include large amounts of food reserves—the yolk and the egg white. The functioning part of the cell is a small mass on the surface of the yolk. Why are most cells so small? If you consider what a cell must do to maintain homeostasis and to grow, it may be easier to understand the reasons for its small size. A cell must take in food and other materials and must rid itself of waste products generated by metabolic reactions. Everything that enters or leaves a cell must pass through its plasma membrane. The plasma membrane contains specialized “pumps” and channels with “gates” that selectively regulate the passage of materials into and out of the cell. The plasma membrane must be large enough relative to the cell volume to keep up with the demands of regulating the passage of materials. Thus, a critical factor in determining cell size is the ratio of its surface area (the plasma membrane) to its volume (FIG. 4-2). As a cell becomes larger, its volume increases at a greater rate than its surface area (its plasma membrane), which effectively places an upper limit on cell size. Above some critical size, the number of molecules required by the cell could not be transported

into the cell fast enough to sustain its needs. In addition, the cell would not be able to regulate its concentration of various ions or efficiently export its wastes. Of course, not all cells are spherical or cuboid. Because of their shapes, some very large cells have relatively favorable ratios of surface area to volume. In fact, some variations in cell shape represent a strategy for increasing the ratio of surface area to volume. For example, many large plant cells are long and thin, which increases their surface area–to-volume ratio. Some cells, such as epithelial cells lining the small intestine, have fingerlike projections of the plasma membrane, called microvilli, that significantly increase the surface area for absorbing nutrients and other materials (see Fig. 47-10). Another reason for the small size of cells is that, once inside, molecules must be transported to the locations where they are converted into other forms. Because cells are small, the distances molecules travel within them are relatively short. Thus, molecules are rapidly available for cell activities.

Cell size and shape are adapted to function The sizes and shapes of cells are adapted to the particular functions they perform. Some cells, such as amoebas and white blood

1 mm 2 mm 2 mm Surface Area (mm2)

Surface area = height × width × number of sides × number of cubes

Volume (mm3)

Volume = height × width × length × number of cubes

Surface Area/ Volume Ratio

Surface area/ volume

1 mm

24

48

(2 × 2 × 6 × 1)

(1 × 1 × 6 × 8)

8

8

(2 × 2 × 2 × 1)

(1 × 1 × 1 × 8)

3 (24: 8)

6 (48 : 8)

FIGURE 4-2 Surface area–to-volume ratio The surface area of a cell must be large enough relative to its volume to allow adequate exchange of materials with the environment. Although their volumes are the same, eight small cells have a much greater surface area (plasma membrane) in relation to their total volume than one large cell does. In the example shown, the ratio of the total surface area to total volume of eight 1 mm cubes is double the surface area–to-volume ratio of the single large cube.

cells, change their shape as they move about. Sperm cells have long, whiplike tails, called flagella, for locomotion. Nerve cells have long, thin extensions that enable them to transmit messages over great distances. The extensions of some nerve cells in the human body may be as long as 1 meter! Certain epithelial cells are almost rectangular and are stacked much like building blocks to form sheetlike tissues. (Epithelial tissue covers the body and lines body cavities.)

Review ■

■ ■

How does the cell theory contribute to our understanding of the evolution of life? How does the plasma membrane help maintain homeostasis? Why is the relationship between surface area and volume of a cell important in detemining cell-size limits?

4.2 METHODS FOR STUDYING CELLS ■ ■ LEARNING OBJECTIVE 4 Describe methods that biologists use to study cells, including microscopy and cell fractionation.

One of the most important tools biologists use for studying cell structures is the microscope. Using a microscope he had made, Robert Hooke, an English scientist, first described cells in 1665 in his book Micrographia. Hooke examined a piece of cork and drew and described what he saw. Hooke chose the term cell because the tissue reminded him of the small rooms monks lived in. Interestingly, what

Hooke saw were not actually living cells but the walls of dead cork cells (FIG. 4-3a). Much later, scientists recognized that the interior enclosed by the walls is the important part of living cells. A few years after Hooke’s discovery and inspired by Hooke’s work, the Dutch naturalist Antonie van Leeuwenhoek viewed living cells with small lenses that he made. Leeuwenhoek was highly skilled at grinding lenses and was able to magnify images more than 200 times. Among his important discoveries were bacteria, protists, blood cells, and sperm cells. Leeuwenhoek was among the first scientists to report cells in animals. Leeuwenhoek was a merchant and not formally trained as a scientist. However, his skill, curiosity, and diligence in sharing his discoveries with scientists at the Royal Society of London brought an awareness of microscopic life to the scientific world. Unfortunately, Leeuwenhoek did not share his techniques, and it was more than 100 years later, in the late 19th century, before microscopes were sufficiently developed for biologists to seriously focus their attention on the study of cells.

Light microscopes are used to study stained or living cells The light microscope (LM), the type used by most students, consists of a tube with glass lenses at each end. Because it contains several lenses, the modern light microscope is referred to as a compound microscope. Visible light passes through the specimen being observed and through the lenses. Light is refracted (bent) by the lenses, magnifying the image. Images obtained with light microscopes are referred to as light micrographs, or LMs. Two features of a microscope determine how clearly a small object can be viewed: magnification and resolving power. Magnification is the ratio of the size of the image seen with the microscope to the actual size of the object. The best light microscopes usually magnify an object no more than 2000 times. Resolution, or resolving power, is the capacity to distinguish fine detail in an image; it is defined as the minimum distance between two points at which they can both be seen separately rather than as a single, blurred point. Resolving power depends on the quality of the lenses and the wavelength of the illuminating light. As the wavelength decreases, the resolution increases. The visible light used by light microscopes has wavelengths ranging from about 400 nm (violet) to 700 nm (red); this limits the resolution of the light microscope to details no smaller than the diameter of a small bacterial cell (about 0.2 μm). By the early 20th century, refined versions of the light microscope became available. The interior of many cells is transparent and it is difficult to discern specific cell structures. Organic chemists have contributed greatly to light microscopy by developing biological stains that enhance contrast in the microscopic image. Staining has enabled biologists to discover the many different internal cell structures,

RESEARCH METHOD

(e) Differential-interference-contrast (Nomarski).

100 μm

(c) Dark-field.

100 μm

Dennis Kunkel/Visuals Unlimited

100 μm

M. I. Walker/Photo Researchers

100 μm

Michael Abbey/Photo Researchers

How Is It Done?

100 μm

From Hooke’s Micrographia 1665

(b) Bright-field (stained).

(a) Robert Hooke's drawing of cork cells.

(d) Phase contrast.

Wim van Egmond/Visuals Unlimited

Cells are too small to be studied with the naked eye. Biologists use microscopes to view cells and structures inside cells. Many types of microscopes have been developed. Here we look at photomicrographs of Paramecia (a group of ciliated protists) made using several kinds of light microscopes.

Wim van Egmond/Visuals Unlimited

Why Is It Used?

(f) Confocal. Plasma membrane and contractile vacuoles are stained red. Proton pumps associated with the contractile vacuoles are stained yellow.

Using a crude microscope that he constructed, Robert Hooke looked at a thin slice of cork and drew what he saw. Biologists now view cells in more detail using the more sophisticated microscopes and techniques that have been developed. In the light microscope, a beam of light passes through the specimen being observed and through the lenses. The lenses refract the light, which magnifies the image. Bright-field microscopy can be enhanced by staining. The phase contrast and differential-interference-contrast microscopes enhance detail by increasing the differences in optical density in different regions of the cells.

FIGURE 4-3 Animated Using light microscopy

the organelles. Unfortunately, most methods used to prepare and stain cells for observation also kill them in the process. Light microscopes with special optical systems now permit biologists to study living cells. In bright-field microscopy, an image is formed by transmitting light through a cell (or other specimen) (FIG. 4-3b). Because there is little contrast, the details of cell structure are not visible. In dark-field microscopy, rays of light are directed

from the side, and only light scattered by the specimen enters the lenses. The cell is seen as a bright image against a dark background (FIG. 4-3c). The specimen does not need to be stained. Phase contrast microscopy and Nomarski differential-interferencecontrast microscopy take advantage of variations in density within the cell (FIG. 4-3d and e). These variations in density cause differences in the way various regions of the cytoplasm refract (bend)

light. Using these microscopes, scientists can observe living cells in action; they can view numerous internal structures that are constantly changing shape and location. Cell biologists use the fluorescence microscope to detect the locations of specific molecules in cells. In the fluorescence microscope, filters transmit light that is emitted by fluorescently stained molecules. Fluorescent stains (like paints that glow under black light) are molecules that absorb light energy of one wavelength and then release some of that energy as light of a longer wavelength. These stains bind specifically to DNA or to specific protein molecules. The molecules absorb ultraviolet light and emit light of a different color. Cells can be stained, and the location of the labeled molecules can be determined, by observing the source of the fluorescent light within the cell. Biologists commonly use a fluorescent molecule known as green fluorescent protein (GFP) that occurs naturally in jellyfish. GFP has been valuable in observing specific proteins in living cells. Some fluorescent stains chemically bond to antibodies, protein molecules important in internal defense. The antibody binds to a highly specific region of a molecule in the cell. A single type of antibody molecule binds to only one type of structure, such as a part of a specific protein or some of the sugars in a specific polysaccharide. Purified fluorescent antibodies known to bind to a specific protein are used to determine where that protein is located within the cell. Confocal microscopy produces a sharper image than standard fluorescence microscopy (FIG. 4-3f). Living cells that have been labeled with fluorescent dye are mounted on a microscope slide. The confocal laser scanning microscope is a computerized microscope. A beam of ultraviolet light from a laser is focused at a specific depth within the cell. The fluorescent label emits visible light, and the investigator can visualize objects in a single plane of focus. The microscope produces optical sections (see the photomicrograph in the chapter introduction). A computer reassembles the images so that a series of optical sections from different planes of the cell can be used to construct a three-dimensional image. Powerful computer-imaging methods greatly improve the resolution of structures labeled by fluorescent dyes.

Electron microscopes provide a high-resolution image that can be greatly magnified Even with improved microscopes and techniques for staining cells, ordinary light microscopes can distinguish only the gross details of many cell parts (FIG. 4-4a). In many cases, you can clearly see only the outline of an organelle. With the development of the electron microscope (EM), which came into wide use in the 1950s, researchers could begin to study the fine details, or ultrastructure, of cells. The resolving power for the average human adult eye is about 100 μm. The best light microscopes have a resolving power of about 0.2 μm (200 nm). In comparison, some electron microscopes have resolving powers of just less than 1 nm. This is possible because electrons have very short wavelengths, on the order of about 0.1 to 0.2 nm. Although such resolution is difficult to achieve with biological material, researchers can approach that resolution when examining isolated molecules such as proteins and DNA. This high degree of resolution permits magnifications of more than 1 million

times as compared to typical magnifications of no more than 1500 to 2000 times in light microscopy. The image formed by the electron microscope is not directly visible. The electron beam itself consists of energized electrons, which, because of their negative charge, can be focused by electromagnets just as images are focused by glass lenses in a light microscope (FIG. 4-4b). Two main types of electron microscopes are the transmission electron microscope (TEM) and the scanning electron microscope (SEM). The acronyms TEM and SEM also identify that a micrograph was prepared using a transmission or scanning EM. Electron micrographs are black and white. They are often colorized to highlight various structures. In transmission electron microscopy, the specimen is embedded in plastic and then cut into extraordinarily thin sections (50 to 100 nm thick) with a glass or diamond knife. A section is then placed on a small metal grid. The electron beam passes through the specimen and then falls onto a photographic plate or a fluorescent screen. When you look at TEMs in this chapter (and elsewhere), keep in mind that each represents only a thin cross section of a cell. Researchers detect certain specific molecules in electron microscope images by using antibody molecules to which very tiny gold particles are bound. The dense gold particles block the electron beam and identify the location of the proteins recognized by the antibodies as precise black spots on the electron micrograph. In the scanning electron microscope, the electron beam does not pass through the specimen. Instead, the specimen is coated with a thin film of gold or some other metal. When the electron beam strikes various points on the surface of the specimen, secondary electrons are emitted whose intensity varies with the contour of the surface. The recorded emission patterns of the secondary electrons give a 3-D picture of the surface (FIG. 4-4c). The SEM provides information about the shape and external features of the specimen that cannot be obtained with the TEM. Note that the LM, TEM, and SEM are focused by similar principles. A beam of light or an electron beam is directed by the condenser lens onto the specimen and is magnified by the objective lens and the eyepiece in the light microscope or by the objective lens and the projector lens in the TEM. The TEM image is focused onto a fluorescent screen, and the SEM image is viewed on a type of television screen. Lenses in electron microscopes are actually electromagnets that bend the beam of electrons. Several other types of microscopes are now available, including two additional kinds of electron microscopes. Digital microscopes that use cameras to send a digital image to a monitor have also been developed.

Biologists use biochemical techniques to study cell components The EM is a powerful tool for studying cell structure, but it has limitations. The methods used to prepare cells for electron microscopy kill them and may alter their structure. Furthermore, electron microscopy provides few clues about the functions of organelles and other cell components. To determine what organelles actually do, researchers use a variety of biochemical techniques. Cell fractionation is a technique for separating (fractionating) different parts of cells so that they can be studied by physical and chemical methods. Generally, cells are broken apart in a

RESEARCH METHOD Why Is It Used?

Electron microscopes have much greater resolving power than light microscopes. Electron microscopes can also magnify images to a much greater extent than light microscopes. Transmission electron microscope

Light microscope

Scanning electron microscope Electron gun

Light beam

Electron beam

Ocular lens

First condenser lens (electromagnet) Specimen

Projector lens (electromagnetic)

Condenser lens

Scanning coil Final (objective) lens

Objective lens Specimen

Second condenser lens

Cathode ray tube synchronized with scanning coil

Secondary electrons

Light source

Specimen Electron detector

Courtesy of T. K. Maugel, University of Maryland

Film or screen

100 μm

(a) A phase contrast light microscope can be used to view stained or living cells, but at relatively low resolution.

How Is It Done?

100 μm

1 μm

(b) The transmission electron microscope (TEM) produces a high-resolution image that can be greatly magnified. A small part of a thin slice through the Paramecium is shown.

(c) The scanning electron microscope (SEM) provides a clear view of surface features.

In an electron microscope, a beam of electrons is focused on or through the specimen. Instead of glass lenses, electromagnetic lenses are used to form the image. Here we compare images of the protist Paramecium made using a phase contrast light microscope with images made using two types of electron microscopes. These three microscopes produce distinctive images of cells.

FIGURE 4-4 Animated Using electron microscopes

blender. The resulting mixture, called the cell homogenate, is subjected to centrifugal force by spinning in a centrifuge (FIG. 4-5a). The powerful ultracentrifuge can spin at speeds exceeding 100,000 revolutions per minute (rpm), generating a centrifugal force of 500,000 × G (a G is equal to the force of gravity). Centrifugal force separates the extract into two fractions: a pellet and a supernatant. The pellet that forms at the bottom of the tube contains heavier materials, such as nuclei, packed together. The supernatant, the liquid above the pellet, contains lighter particles, dissolved molecules, and ions. After the pellet is removed, the supernatant can be centrifuged again at a higher speed to obtain a pellet that contains the next-heaviest cell components, for example, mitochondria and chloroplasts. In

differential centrifugation, the supernatant is spun at successively higher speeds, permitting various cell components to be separated on the basis of their different sizes and densities (FIG. 4-5b). The pellets can be resuspended and their components can be further purified by density gradient centrifugation. In this procedure, the centrifuge tube is filled with a series of solutions of decreasing density. For example, sucrose solutions can be used. The concentration of sucrose is highest at the bottom of the tube and decreases gradually so that it is lowest at the top. The resuspended pellet is placed in a layer on top of the density gradient. Because the densities of organelles differ, each migrates during centrifugation. Each type of organelle forms a band at the position in the gradient where its own density equals that of the sucrose solution (FIG. 4-5c).

RESEARCH METHOD Why Is It Used?

Cell fractionation is used to separate (fractionate) cell components according to their size and density.

Centrifuge rotor Centrifugal force

Centrifugal force

Hinged bucket containing tube

(a) Centrifugation. Due to centrifugal force, large or very dense particles move toward the bottom of a tube and form a pellet.

Centrifuge supernatant 20,000 × G

Centrifuge supernatant 100,000 × G

10 minutes

30 minutes

90 minutes

Disrupt cells in buffered solution Nuclei in pellet

Mitochondria, chloroplasts in pellet

Microsomal pellet (contains ER, Golgi, plasma membrane)

(b) Differential centrifugation.

How Is It Done?

Low sucrose concentration Sucrose density gradient

Centrifuge 600 × G

Resuspended microsomal pellet on top of sucrose gradient

100,000 × G High sucrose concentration

Plasma membrane Golgi ER

(c) Density gradient centrifugation.

Cells are broken up in a blender. The cell homogenate (the resulting mixture) is then spun in a centrifuge. As a result of centrifugal force, the heaviest cell components, the nuclei, form a pellet at the bottom of the tube. The supernatant (the liquid above the pellet) can then be spun at a higher speed. The next heaviest component, the mitochondria and chloroplasts, form a pellet, and the supernatant can be spun at a higher speed. This process can be repeated several times. The pellet can be further purified by density gradient centrifugation (see text for further explanation).

FIGURE 4-5 Cell fractionation

Purified organelles can then be studied. For example, they can be examined to determine what kinds of proteins and other molecules they might contain, or what types of chemical reactions take place within them.

Recall from Chapter 1 that two basic types of cells are known: prokaryotic cells and eukaryotic cells. Bacteria and archaea are prokaryotic cells. All other known organisms consist of eukaryotic cells.

Review

Organelles of prokaryotic cells are not surrounded by membranes

■ ■

What is the main advantage of the electron microscope? Explain. What is cell fractionation? Describe the process.

4.3 PROKARYOTIC AND EUKARYOTIC CELLS ■ ■ LEARNING OBJECTIVES 5 6

Compare and contrast the general characteristics of prokaryotic and eukaryotic cells, and contrast plant and animal cells. Describe three functions of cell membranes.

Prokaryotic cells are typically smaller than eukaryotic cells. In fact, the average prokaryotic cell is only about 1/10 the diameter of the average eukaryotic cell. In prokaryotic cells, the DNA is typically located in a limited region of the cell called a nuclear area, or nucleoid. Unlike the nucleus of eukaryotic cells, the nuclear area is not enclosed by a membrane (FIG. 4-6). The term prokaryotic, meaning “before the nucleus,” refers to this major difference between prokaryotic and eukaryotic cells. Other types of internal membrane–enclosed organelles are also absent in prokaryotic cells. Like eukaryotic cells, prokaryotic cells have a plasma membrane that surrounds the cell. The plasma membrane confines the

contents of the cell to an internal compartment. In some prokaryotic cells, the plasma membrane may be folded inward to form a complex of membranes along which many of the cell’s metabolic reactions take place. Most prokaryotic cells have cell walls, which are extracellular structures that enclose the entire cell, including the plasma membrane. Many prokaryotes have flagella (sing., flagellum), long fibers that project from the surface of the cell. Prokaryotic flagella, which operate like propellers, are important in locomotion. Their structure is different from that of flagella found in eukaryotic cells. Some prokaryotes also have hairlike projections called fimbriae, which are used to adhere to one another or to attach to cell surfaces of other organisms. The dense internal material of the bacterial cell contains ribosomes, small complexes of ribonucleic acid (RNA) and protein that synthesize polypeptides. The ribosomes of prokaryotic cells are smaller than those found in eukaryotic cells. Prokaryotic cells also contain storage granules that hold glycogen, lipid, or phosphate compounds. This chapter focuses primarily on eukaryotic cells. Prokaryotes are discussed in more detail in Chapter 25.

Fimbriae

Storage granule

Flagellum Ribosome Cell wall Plasma membrane DNA

Nuclear area

Dr. Klaus Boller/Science Photo Library/Photo Researchers, Inc.

Capsule

0.5 μm

FIGURE 4-6 Animated Structure of a prokaryotic cell This colorized TEM shows a thin lengthwise slice through an Escherichia coli bacterium. Note the prominent nuclear area containing the genetic material (DNA). E. coli is a normal inhabitant of the human intestine, but under certain conditions some strains can cause infections.

Membranes divide the eukaryotic cell into compartments Eukaryotic cells are characterized by highly organized membraneenclosed organelles, including a prominent nucleus, which contains DNA, the hereditary material. The term eukaryotic means “true nucleus.” Early biologists thought cells consisted of a homogeneous jelly, which they called protoplasm. With the electron microscope and other modern research tools, perception of the environment within the cell has been greatly expanded. We now know that the cell is highly organized and complex (FIGS. 4-7 through 4-10). The eukaryotic cell has its own control center, internal transportation system, power plants, factories for making needed materials, packaging plants, and even a “self-destruct” system. Biologists refer to the part of the cell outside the nucleus as cytoplasm and the part of the cell within the nucleus as nucleoplasm. Various organelles are suspended within the fluid component of the cytoplasm, which is called the cytosol. The term cytoplasm includes both the cytosol and all the organelles other than the nucleus. The many specialized organelles of eukaryotic cells solve some of the problems associated with large size, so eukaryotic cells can be larger than prokaryotic cells. Eukaryotic cells also differ from prokaryotic cells in having a supporting framework, or cytoskeleton, important in maintaining shape and transporting materials within the cell. Some organelles are present only in specific cells. For example, chloroplasts, structures that trap sunlight for energy conversion, are only in cells that carry on photosynthesis, such as certain plant or algal cells. Cells of fungi and plants are surrounded by a cell wall external to the plasma membrane. Plant cells also contain a large, membrane-enclosed vacuole. We discuss these and other differences among major types of cells throughout this chapter. Plant and animal cells are compared in Figures 4-7 and 4-8 and also in Figures 4-9 and 4-10. Cell membranes have unique properties that enable membranous organelles to carry out a wide variety of functions. For example, cell membranes never have free ends. As a result, a membranous organelle always contains at least one enclosed internal space or compartment. These membrane-enclosed compartments allow certain cell activities to be localized within specific regions of the cell. Reactants located in only a small part of the total cell volume are far more likely to come in contact, dramatically increasing the rate of the reaction. On the other hand, membrane-enclosed compartments keep certain reactive compounds away from other parts of the cell that they might adversely affect. Compartmentalizing also allows many different activities to go on simultaneously. Membranes serve as important work surfaces. For example, many chemical reactions in cells are carried out by enzymes that are bound to membranes. Because the enzymes that carry out successive steps of a series of reactions are organized close together on a membrane surface, certain series of chemical reactions occur more rapidly. Membranes allow cells to store energy. The membrane serves as a barrier that is somewhat analogous to a dam on a river. As we will discuss in Chapter 5, there is both an electric charge difference and a concentration difference on the two sides of the membrane.

the endomembrane system because they function somewhat independently of other membranous organelles.) Some organelles have direct connections between their membranes and other compartments. Others transport materials in vesicles, small, membrane-enclosed sacs formed by “budding” from

Courtesy of Dr. Kenneth Miller, Brown University

Dr. Susumu Ito, Harvard Medical School

These differences constitute an electrochemical gradient. Such gradients store energy and so have potential energy (discussed in Chapter 7). As particles of a substance move across the membrane from the side of higher concentration to the side of lower concentration, the cell can convert some of this potential energy to the chemical energy of ATP molecules. This process of energy conversion (discussed in Chapters 7, 8, and 9) is a basic mechanism that cells use to capture and convert the energy necessary to sustain life. In a eukaryotic cell, several types of membranes make up the internal membrane system, or endomembrane system. In Figures 4-7 and 4-8 (also see Figs. 4-9 and 4-10), notice how membranes divide the cell into many compartments: the nucleus, endoplasmic reticulum (ER), Golgi complex, lysosomes, vesicles, and vacuoles. Although it is not internal, the plasma membrane is also included because it participates in the activities of the endomembrane system. (Mitochondria and chloroplasts are also separate compartments but are not generally considered part of

5 μm Chromatin

Plasma membrane

Nucleolus

5 μm Nucleus

Starch grain

Golgi complex

Rough endoplasmic reticulum Ribosomes

Vacuole

Plasma membrane

Prolamellar body

Intercellular space

Nucleus

Golgi complex

Ribosomes

Rough endoplasmic reticulum

Zymogen granules

Mitochondria Smooth endoplasmic reticulum

FIGURE 4-8 TEM of a human pancreas cell and an interpretive drawing

Chloroplasts

Cell wall

FIGURE 4-7 TEM of a plant cell and an interpretive drawing Most of this cross section of a cell from the leaf of a young bean plant (Phaseolus vulgaris) is dominated by a vacuole. Prolamellar bodies are membranous regions typically seen in developing chloroplasts. Can you label the TEM?

Most of the structures of a typical animal cell are present. However, like most cells, this one has certain structures associated with its specialized functions. Pancreas cells such as the one shown here secrete large amounts of digestive enzymes. The large, dark, circular bodies in the TEM and the corresponding structures in the drawing are zymogen granules, which contain inactive enzymes. When released from the cell, the enzymes catalyze chemical reactions such as the breakdown of peptide bonds of ingested proteins in the intestine. Most of the membranes visible in this section are part of the rough endoplasmic reticulum, an organelle specialized to manufacture protein. Can you label the TEM?

Bloom and Fawcett Textbook of Histology

Cristae

Bloom and Fawcett Textbook of Histology

Membranous sacs

Golgi complex

Mitochondrion

Cell wall Plasma membrane

Granum

Stroma

Chloroplast

Bloom and Fawcett Textbook of Histology

E. H. Newcomb and W. P. Wergin, Biological Photo Service

Smooth ER

Dr. Donald Fawcett/Visuals Unlimited, Inc.

Vacuole

Nuclear envelope Nucleolus Nuclear pores

Rough ER

Chromatin

Ribosomes Rough and smooth endoplasmic reticulum (ER)

FIGURE 4-9 Animated Composite diagram of a plant cell Plant cells typically have a cell wall, chloroplasts, and prominent vacuoles. The TEMs show specific structures or areas of the cell. Some plant cells do not have all the organelles shown here. For example, leaf and stem cells that carry on photosynthesis contain chloroplasts, whereas root cells do not. Many of the organelles, such as the nucleus, mitochondria, and endoplasmic reticulum (ER), are characteristic of all eukaryotic cells.

Nucleus

Chromatin

Nucleolus

Bloom and Fawcett Textbook of Histology

Nuclear pores

Membranous sacs of Golgi

Bloom and Fawcett Textbook of Histology

Nuclear envelope

Golgi complex

Nucleus

Plasma membrane

Lysosome Nuclear envelope

Cristae

Dr. Donald Fawcett/Visuals Unlimited, Inc.

B. F. King, Biological Photo Service

Rough ER

Bloom and Fawcett Textbook of Histology

Ribosomes

Smooth ER Rough and smooth endoplastic reticulum (ER)

Centrioles

FIGURE 4-10 Animated Composite diagram of an animal cell This generalized animal cell is shown in a realistic context surrounded by adjacent cells, which cause it to be slightly compressed. The TEMs show the structure of various organelles. Depending on the cell type, certain organelles may be more or less prominent.

Mitochondrion

the membrane of another organelle. Vesicles carry cargo from one organelle to another. A vesicle can form as a “bud” from the membrane of one organelle and then move to another organelle to which it fuses, thus delivering its contents into another compartment.

■ ■ ■

EXPERIMENT 1 QUESTION: What controls the shape of the cap in Acetabularia? HYPOTHESIS: Something in the stalk or holdfast of Acetabularia controls the shape of the cap.

Review ■

KEY EXPERIMENTS

What are two important differences between prokaryotic and eukaryotic cells? What are three ways that a plant cell might differ from an animal cell? How do membrane-enclosed organelles facilitate cell metabolism? What organelles belong to the endomembrane system?

EXPERIMENT: Hämmerling removed the caps from A. mediterranea and A. crenulata. He then grafted together the two capless algae (FIG. a).

4.4 THE CELL NUCLEUS Nucleus

■ ■ LEARNING OBJECTIVE 7

Cap Stalk Holdfast

Describe the structure and functions of the nucleus.

L. Sims/Visuals Unlimited

Typically, the nucleus is the most prominent organelle in the cell. It is usually spherical or oval in shape and averages 5 μm in diameter. Because of its size and the fact that it often occupies a relatively fixed position near the center of the cell, some early investigators guessed long before experimental evidence was available that the nucleus served as the control center of the cell. In the 19th century, biologists discovered that the marine alga Acetabularia consists of a single cell (FIG. 4-11). At up to 5 cm (2 in) in length, Acetabularia is small for a seaweed but gigantic for a cell. It consists of a rootlike holdfast; a long, cylindrical stalk; and a cuplike cap. The nucleus is in the holdfast, about as far away from the cap as it can be. Because it is a single giant cell, Acetabularia is easy for researchers to manipulate. If the cap of Acetabularia is removed experimentally, another one grows after a few weeks. Such regeneration is common among

FIGURE 4-11 LM of Acetabularia To the romantically inclined, the little seaweed Acetabularia resembles a mermaid’s wineglass, although the literal translation of its name, “vinegar cup,” is somewhat less elegant. Acetabularia, which consists of a single cell, has been a model organism for investigating the role of the nucleus.

A. mediterranea A. crenulata

(a)

RESULTS AND CONCLUSION: The algae regenerated a common cap with characteristics intermediate between those of the two species involved. This experiment demonstrated that something in the stalk or holdfast controls cap shape.

FIGURE 4-12 Acetabularia and the control of cell

activities Biologist J. Hämmerling used two species of the single-celled seaweed Acetabularia for most of his experiments: A. mediterranea, which has a smooth cap, and A. crenulata, which has a cap divided into a series of fingerlike projections. When the cap is removed, the kind of cap that is regenerated depends on the species of Acetabularia used in the experiment. As you might expect, A. crenulata regenerates a “cren” cap, and A. mediterranea regenerates a “med” cap.

simple organisms. This fact attracted the attention of investigators, especially Danish biologist J. Hämmerling, who became interested in whether a relationship exists between the nucleus and the physical characteristics of the alga. Hämmerling (during the 1930s to 1950s) performed brilliant experiments that in many ways laid the foundation for much of our modern knowledge of the nucleus. Some of his experiments are summarized in FIGURE 4-12. Cell biologists extended these early findings as they developed our modern view of information flow and control in the cell. We now know that the cell stores information in the form of DNA, and most of the cell’s DNA is located inside the nucleus. The nuclear envelope consists of two concentric membranes that separate the nuclear contents from the surrounding cytoplasm (FIG. 4-13). These membranes are separated by about 20 to 40 nm. At intervals the membranes come together to form nuclear pores. Each nuclear pore consists of a molecular complex made up of many copies of about 30 different proteins. Nuclear pores regulate the passage of materials between nucleoplasm and cytoplasm. A fibrous network of protein filaments, called the nuclear lamina, forms an inner lining for the nuclear envelope. The nuclear

EXPERIMENT 2 QUESTION: Does the stalk or the holdfast control the shape of the cap in Acetabularia? HYPOTHESIS: Something in the holdfast of Acetabularia controls the shape of the cap. EXPERIMENT: Hämmerling removed the caps from A. mediterranea and A. crenulata. Then he severed the stalks from the holdfasts. By telescoping the cell walls of the two species into each other, Hämmerling was able to attach a section of the stalk of one species to a holdfast of the other species (see FIG. b).

Caps removed

Stalks and holdfasts exchanged

(b)

Caps removed again

First regenerated caps

Second regenerated caps

RESULTS AND CONCLUSION: The results were surprising. The caps that regenerated were characteristic not of the species donating the holdfasts but of those donating the stalks! However, when the caps were removed once again, this time the caps that regenerated were characteristic of the species that donated the holdfasts. This continued to be the case no matter how many more times the regenerated caps were removed. From these results Hämmerling deduced that the ultimate control of the Acetabularia cell is associated with the holdfast. Because there is a time lag before the holdfast appears to take over, he hypothesized it produces some temporary cytoplasmic messenger substance whereby it exerts its control. Hämmerling further hypothesized that the grafted stalks initially contain enough of the substance from their former holdfasts to regenerate a cap of the former shape. But this still left the question of how the holdfast exerts its apparent control. An obvious suspect was the nucleus. EXPERIMENT 3 QUESTION: Does the nucleus in the holdfast control the shape of the cap in Acetabularia? HYPOTHESIS: The nucleus in the holdfast of Acetabularia controls the shape of the cap. EXPERIMENT: Hämmerling removed the nucleus and cut off the cap of Acetabularia (FIG. c). A new cap typical of the species regenerated. Acetabularia, however, can usually regenerate only once without a nucleus. The researcher inserted a nucleus of another species and cut the cap off once again (FIG. d).

Eventually

(c)

The characteristics of the cell are governed by the messenger substance and, therefore, ultimately by the nucleus.

Eventually

(d)

Messenger substance

(e)

The nucleus produces the messenger.

RESULTS AND CONCLUSION: A new cap regenerated that was characteristic of the species of the nucleus. When two kinds of nuclei were inserted, the regenerated cap was intermediate in shape between those of the species that donated the nuclei. As a result of these and other experiments, biologists began to develop some basic ideas about the control of cell activities. The holdfast controls the cell because the nucleus is located there. Further, the nucleus is the apparent source of some “messenger substance” that temporarily exerts control but is limited in quantity and cannot be produced without the nucleus (FIG. e). This information helped provide a starting point for research on the role of nucleic acids in the control of all cells. Much later, the “messenger substance” was characterized and named messenger RNA (mRNA).

KEY POINT

The nucleus contains DNA and is the control center of the cell.

Chromatin Nucleolus

Nuclear pores

Nuclear pore

Nuclear envelope

R. Kessel and G. Shih/Visuals Unlimited

Rough ER

0.25 μm

(b)

ER continuous with outer membrane of nuclear envelope

Bloom and Fawcett Textbook of Histology

Nucleoplasm

Outer nuclear envelope

Nuclear pore

2 μm

(a)

(c)

Nuclear pore proteins

Inner nuclear envelope

FIGURE 4-13 Animated The cell nucleus (a) The TEM and interpretive drawing show that the nuclear envelope, composed of two concentric membranes, is perforated by nuclear pores (red arrows). The outer membrane of the nuclear envelope is continuous with the membrane of the ER (endoplasmic reticulum). The nucleolus

lamina supports the inner nuclear membrane and helps organize the nuclear contents. It also plays a role in DNA duplication and in regulating the cell cycle. Mutations in genes encoding proteins that make up the nuclear lamina are associated with several human genetic diseases, including muscular dystrophies and premature aging (progeria). When a cell divides, the information stored in DNA must be reproduced and passed intact to the two daughter cells. DNA has the unique ability to make an exact duplicate of itself through a process called replication. DNA molecules include sequences of nucleotides called genes, which contain the chemically coded instructions for producing the proteins needed by the cell. The

is not bounded by a membrane. (b) TEM of nuclear pores. A technique known as freeze-fracture was used to split the membrane. (c) The nuclear pores, which are made up of proteins, form channels between the nucleoplasm and cytoplasm.

nucleus controls protein synthesis by transcribing its information in messenger RNA (mRNA) molecules. Messenger RNA moves into the cytoplasm, where proteins are manufactured. DNA is associated with RNA and certain proteins, forming a complex known as chromatin. This complex appears as a network of granules and strands in cells that are not dividing. Although chromatin appears disorganized, it is not. Because DNA molecules are extremely long and thin, they must be packed inside the nucleus in a regular fashion as part of structures called chromosomes. In dividing cells, the chromosomes become visible as distinct threadlike structures. If the DNA in the 46 chromosomes of one human cell could be stretched end to end, it would extend for 2 meters!

Most nuclei have one or more compact structures called nucleoli (sing., nucleolus). A nucleolus, which is not enclosed by a membrane, usually stains differently than the surrounding chromatin. Each nucleolus contains a nucleolar organizer, made up of chromosomal regions containing instructions for making the type of RNA in ribosomes. This ribosomal RNA (rRNA) is synthesized in the nucleolus. The proteins needed to make ribosomes are synthesized in the cytoplasm and imported into the nucleolus. Ribosomal RNA and proteins are then assembled into ribosomal subunits that leave the nucleus through the nuclear pores.

Review ■ ■

How does the nucleus store information? What is the function of the nuclear envelope?

4.5 ORGANELLES IN THE CYTOPLASM ■ ■ LEARNING OBJECTIVES 8 Distinguish between smooth and rough endoplasmic reticulum in terms of both structure and function.

9 Trace the path of proteins synthesized in the rough endoplasmic reticulum as they are processed, modified, and sorted by the Golgi complex and then transported to specific destinations. 10 Describe the functions of lysosomes, vacuoles, and peroxisomes. 11 Compare the functions of mitochondria and chloroplasts, and discuss ATP synthesis by each of these organelles.

(ER), forms a network that makes up a significant part of the total volume of the cytoplasm in many cells. A higher-magnification TEM of the ER is shown in FIGURE 4-14. Remember that a TEM represents only a thin cross section of the cell, so there is a tendency to interpret the ER as a series of tubes. In fact, many ER membranes consist of a series of tightly packed and flattened, saclike structures that form interconnected compartments within the cytoplasm. The internal space the membranes enclose is called the ER lumen. In most cells the ER lumen forms a single internal compartment that is continuous with the compartment formed between the outer and inner membranes of the nuclear envelope (see Fig. 4-13). The membranes of other organelles are not directly connected to the ER; they form distinct and separate compartments within the cytoplasm. The ER membranes and lumen contain enzymes that catalyze many types of chemical reactions. In some cases the membranes serve as a framework for enzymes that carry out sequential biochemical reactions. The two surfaces of the membrane contain different sets of enzymes and represent regions of the cell with different synthetic capabilities, just as different regions of a factory make different parts of a particular product. Still other enzymes are located within the ER lumen. Two distinct regions of the ER can be distinguished in TEMs: rough ER and smooth ER. Although these regions have different functions, their membranes are connected and their internal spaces are continuous.

The endoplasmic reticulum is a network of internal membranes

Smooth ER synthesizes lipids Smooth ER has a tubular appearance, and its outer membrane surfaces appear smooth. Enzymes in the membranes of the smooth ER catalyze the synthesis of many lipids and carbohydrates. The smooth ER is the primary site for the synthesis of phospholipids and cholesterol needed to make cell membranes. Smooth ER synthesizes steroid hormones, including reproductive hormones, from cholesterol. In liver cells, smooth ER is important in enzymatically breaking down stored glycogen. (The liver helps regulate the concentration of glucose in the blood.) Smooth ER also stores calcium ions. Whereas smooth ER may be a minor membrane component in some cells, extensive amounts of smooth ER are present in others. For example, extensive smooth ER is present in human liver cells, where it synthesizes and processes cholesterol and other lipids and serves as a major detoxification site. Enzymes located along the smooth ER of liver cells break down toxic chemicals such as carcinogens (cancer-causing agents) and many drugs, including alcohol, amphetamines, and barbiturates. The cell then converts these compounds to water-soluble products that it excretes. Interestingly, alcohol and many other drugs stimulate liver cells to produce additional smooth ER, increasing the rate that these cells can detoxify the drugs. Alcohol abuse causes liver inflammation that can lead to cirrhosis and eventual liver failure.

One of the most prominent features in the electron micrographs in Figures 4-9 and 4-10 is a maze of parallel internal membranes that encircle the nucleus and extend into many regions of the cytoplasm. This complex of membranes, the endoplasmic reticulum

Rough ER is important in protein synthesis The outer surface of the rough ER is studded with ribosomes that appear as dark granules. Notice in Figure 4-14 that the lumen side

Cell biologists have identified many types of organelles in the cytoplasm of eukaryotic cells. Among them are the ribosomes, endoplasmic reticulum, Golgi complex, lysosomes, peroxisomes, vacuoles, mitochondria, and chloroplasts. TABLE 4-1 summarizes eukaryotic cell structures and functions.

Ribosomes manufacture proteins Ribosomes are tiny particles found free in the cytoplasm or attached to certain membranes. They consist of RNA and proteins and are synthesized by the nucleolus. Ribosomes contain the enzyme necessary to form peptide bonds, which join amino acids to produce polypeptides (see Chapter 3). Each ribosome has two main components: a large subunit and a small subunit. When the two ribosome subunits join, they function as manufacturing plants that assemble polypeptides. Cells that actively produce a lot of proteins may have millions of ribosomes, and the cell can change the number of ribosomes present to meet its metabolic needs. We will discuss much more about ribosomes in Chapter 13.

TABLE 4-1 Structure Cell Nucleus Nucleus Nucleolus Chromosomes

Eukaryotic Cell Structures and Their Functions Description

Function

Large structure surrounded by double membrane; contains nucleolus and chromosomes Granular body within nucleus; consists of RNA and protein Composed of chromatin, a complex of DNA and protein; condense during cell division, becoming visible as rodlike structures

Information in DNA is transcribed in RNA synthesis; specifies cell proteins Site of ribosomal RNA synthesis; ribosome subunit assembly Contain genes (units of hereditary information) that govern structure and activity of cell

Cytoplasmic Organelles Plasma Membrane boundary of cell membrane Ribosomes Endoplasmic reticulum (ER) Smooth Rough Golgi complex

Granules composed of RNA and protein; some attached to ER, some free in cytosol Network of internal membranes extending through cytoplasm Lacks ribosomes on outer surface Ribosomes stud outer surface Stacks of flattened membrane sacs

Lysosomes

Membranous sacs (in animals)

Vacuoles Peroxisomes

Membranous sacs (mostly in plants, fungi, algae) Membranous sacs containing a variety of enzymes

Mitochondria

Sacs consisting of two membranes; inner membrane is folded to form cristae and encloses matrix Double-membrane structure enclosing internal thylakoid membrane; chloroplasts contain chlorophyll in thylakoid membrane

Plastids (e.g., chloroplasts)

Cytoskeleton Microtubules

Hollow tubes made of subunits of tubulin protein

Microfilaments

Solid, rodlike structures consisting of actin protein

Intermediate filaments Centrioles

Tough fibers made of protein

Cilia

Flagella

Pair of hollow cylinders located near nucleus; each centriole consists of nine microtubule triplets (9 × 3 structure) Relatively short projections extending from surface of cell; covered by plasma membrane; made of two central and nine pairs of peripheral microtubules (9 + 2 structure) Long projections made of two central and nine pairs of peripheral microtubules (9 +2 structure); extend from surface of cell; covered by plasma membrane

of the rough ER appears bare, whereas the outer surface (the cytosolic side) looks rough. The ribosomes attached to the rough ER are known as bound ribosomes; free ribosomes are suspended in the cytosol. The rough ER plays a central role in the synthesis and assembly of proteins. Many proteins that are exported from the cell (such as digestive enzymes), and proteins destined for other organelles, are synthesized on ribosomes bound to the ER membrane. The ri-

Encloses cell contents; regulates movement of materials in and out of cell; helps maintain cell shape; communicates with other cells (also present in prokaryotes) Synthesize polypeptides in both prokaryotes and eukaryotes Synthesizes lipids and modifies many proteins; origin of intracellular transport vesicles that carry proteins Lipid synthesis; drug detoxification; calcium ion storage Manufactures proteins Modifies proteins; packages secreted proteins; sorts other proteins to vacuoles and other organelles Contain enzymes that break down ingested materials; break down damaged or unneeded organelles and proteins Store materials, wastes, water; maintain hydrostatic pressure Site of many diverse metabolic reactions; e.g., break down fatty acids Site of most reactions of cellular respiration; transformation of energy originating from glucose or lipids into ATP energy Chloroplasts are site of photosynthesis; chlorophyll captures light energy; ATP and other energy-rich compounds are produced and then used to convert CO2 to carbohydrate

Provide structural support; have role in cell and organelle movement and cell division; components of cilia, flagella, centrioles, basal bodies Provide structural support; play role in cell and organelle movement and cell division Help strengthen cytoskeleton; stabilize cell shape Mitotic spindle forms between centrioles during animal cell division; may anchor and organize microtubule formation in animal cells; absent in most plant cells Movement of some unicellular organisms; used to move materials on surface of some tissues; important in cell signaling

Cell locomotion by sperm cells and some unicellular organisms

bosome forms a tight seal with the ER membrane. A tunnel within the ribosome connects to an ER pore. Polypeptides are transported through the tunnel and the pore in the ER membrane into the ER lumen. In the ER lumen, proteins are assembled and may be modified by enzymes that add carbohydrates or lipids to them. Other enzymes, called molecular chaperones, in the ER lumen catalyze the efficient folding of proteins into proper conformations. Pro-

ER lumen Mitochondrion cis face

Bloom and Fawcett Textbook of Histology

Ribosomes

trans face

Rough ER

Golgi complex 0.5 μm

Dr. Donald Fawcett/Visuals Unlimited, Inc.

FIGURE 4-15 TEM and an interpretive drawing of the Golgi complex

1 μm

Smooth ER

FIGURE 4-14 Animated Endoplasmic reticulum (ER) The TEM shows both rough and smooth ER in a liver cell.

teins that are not processed correctly, for example, proteins that are misfolded, are transported to the cytosol. There, they are degraded by proteasomes, protein complexes in the cytosol that direct the destruction of defective proteins. Properly processed proteins are transferred to other compartments within the cell by small transport vesicles, which bud off the ER membrane and then fuse with the membrane of some target organelle.

The Golgi complex processes, sorts, and modifies proteins The Golgi complex (also known as the Golgi body or Golgi apparatus) was first described in 1898 by the Italian microscopist Camillo Golgi, who found a way to specifically stain this organelle. However, many investigators thought the Golgi complex was an arti-

fact, and its legitimacy as a cell organelle was not confirmed until cells were studied with the electron microscope in the 1950s. In many cells, the Golgi complex consists of stacks of flattened membranous sacs called cisternae (sing., cisterna). Each cisterna has an internal space, or lumen (FIG. 4-15). In certain regions, cisternae may be distended because they are filled with cell products. The Golgi complex contains a number of separate compartments, as well as some that are interconnected. Each Golgi stack has three areas, referred to as the cis face, the trans face, and a medial region in between. Typically, the cis face (the entry surface) is located nearest the nucleus and receives materials from transport vesicles bringing molecules from the ER. The trans face (the exit surface) is closest to the plasma membrane. It packages molecules in vesicles and transports them out of the Golgi. In a cross-sectional view like that in the TEM in Figure 4-15, many ends of the sheetlike layers of Golgi membranes are distended, an arrangement characteristic of well-developed Golgi complexes in many cells. In some animal cells, the Golgi complex lies at one side of the nucleus; other animal cells and plant cells have many Golgi complexes dispersed throughout the cell. Cells that secrete large amounts of glycoproteins have many Golgi stacks. (Recall from Chapter 3 that a glycoprotein is a protein with a covalently attached carbohydrate.) Golgi complexes of plant cells produce extracellular polysaccharides that are used as components of the cell wall. In animal cells, the Golgi complex manufactures lysosomes. The Golgi complex processes, sorts, and modifies proteins. Researchers have studied the function of the ER, Golgi complex,

KEY POINT

After proteins are synthesized, they are transported through a series of compartments where they are successively modified.

1 Polypeptides synthesized

on ribosomes are inserted into ER lumen. 2 Sugars are added, forming

glycoproteins. 3 Transport vesicles deliver

glycoproteins to cis face of Golgi.

Ribosomes 1

Rough ER Glycoprotein

2 cis face

4 Glycoproteins modified

3

further in Golgi. 4

5 Glycoproteins move to trans

face where they are packaged in transport vesicles.

5

trans face 6

6 Glycoproteins transported to

plasma membrane (or other organelle).

Golgi complex

7 Contents of transport vesicle

released from cell.

Plasma membrane

7

FIGURE 4-16 Animated Protein transport within the cell Glycoproteins are transported from ribosomes into the ER. They are then transported to the Golgi complex, where they are modified. This diagram shows the passage of glycoproteins through compartments of

and other organelles by radioactively labeling newly manufactured molecules and observing their movement through the cell. The general pathway is from ribosomes to the lumen of the rough ER to the Golgi complex and then to some final destination (FIG. 4-16). Proteins that are accurately assembled are transported from the rough ER to the cis face of the Golgi complex in small transport vesicles formed from the ER membrane. Transport vesicles fuse with one another to form clusters that move along microtubules (part of the cytoskeleton) to the Golgi complex. What happens to glycoprotein molecules released into the Golgi complex? One hypothesis holds that the glycoproteins are enclosed in new vesicles that shuttle them from one compartment to another within the Golgi complex. A competing hypothesis postulates that the cisternae themselves may move from cis to trans positions. Both hypotheses may be correct: glycoproteins may be transported by both methods. Regardless of how proteins are moved through the Golgi complex, while there they are modified in different ways, resulting in the formation of complex biological molecules. For example, the carbohydrate part of a glycoprotein (first added to proteins in the

the endomembrane system of a mucus-secreting goblet cell from the lining of the intestine. Mucus consists of a complex mixture of covalently linked proteins and carbohydrates.

rough ER) may be modified. In some cases the carbohydrate component may be a “sorting signal,” a cellular zip code that tags the protein, routing it to a specific organelle. Glycoproteins are packaged in transport vesicles in the trans face. These vesicles pinch off from the Golgi membrane and transport their contents to a specific destination. Vesicles transporting products for export from the cell fuse with the plasma membrane. The vesicle membrane becomes part of the plasma membrane, and the glycoproteins are secreted from the cell. Other vesicles may store glycoproteins for secretion at a later time, and still others are routed to various organelles of the endomembrane system. In summary, here is a typical sequence followed by a glycoprotein destined for secretion from the cell: polypeptides synthesized on ribosomes ¡ protein assembled and carbohydrate component added in lumen of ER ¡ transport vesicles move glycoprotein to Golgi (cis face) ¡ glycoprotein further modified in Golgi ¡ in trans face, glycoproteins packaged in transport vesicles ¡ glycoproteins transported to plasma membrane ¡ contents released from cell

Lysosomes are small sacs of digestive enzymes dispersed in the cytoplasm of most animal cells (FIG. 4-17). Researchers have identified about 40 different digestive enzymes in lysosomes. Because lysosomal enzymes are active under rather acidic conditions, the lysosome maintains a pH of about 5 in its interior. The powerful enzymes and low pH that the lysosome maintains provide an excellent example of the importance of separating functions within the cell into different compartments. Under most normal conditions, the lysosome membrane confines its enzymes and their actions. However, some forms of tissue damage are related to “leaky” lysosomes. Primary lysosomes are formed by budding from the Golgi complex. Their hydrolytic enzymes are synthesized in the rough ER. As these enzymes pass through the lumen of the ER, sugars attach to each molecule, identifying it as bound for a lysosome. This signal permits the Golgi complex to sort the enzyme to the lysosomes rather than to export it from the cell. Bacteria (or debris) engulfed by scavenger cells are enclosed in a vesicle formed from part of the plasma membrane. One or more primary lysosomes fuse with the vesicle containing the ingested material, forming a larger vesicle called a secondary lysosome. Powerful enzymes in the secondary lysosome come in contact with the ingested molecules and degrade them into their components. Under some conditions, lysosomes break down organelles and allow their components to be recycled or used as an energy source. In certain genetic diseases of humans, known as lysosomal storage diseases, one of the digestive enzymes normally present in lysosomes is absent. Its substrate (a substance the enzyme would normally break down) accumulates in the cell, which eventually interferes with cell activities. An example is Tay-Sachs disease, an inherited disease in which a normal lipid cannot be broken down (discussed in Chapter 16). The accumulation of this lipid in brain cells causes mental retardation, blindness, and death before age 4.

Vacuoles are large, fluid-filled sacs with a variety of functions Although lysosomes have been identified in almost all kinds of animal cells, biologists have not identified them in plant and fungal cells. Many functions carried out in animal cells by lysosomes are performed in plant and fungal cells by a large, single, membraneenclosed sac called a vacuole. The membrane of the vacuole, part of the endomembrane system, is called the tonoplast. The term vacuole, which means “empty,” refers to the fact that these organelles have no internal structure. Although some biologists use the terms vacuole and vesicle interchangeably, vacuoles are usually larger structures, sometimes produced by the merging of many vesicles. Vacuoles play a significant role in plant growth and development. Immature plant cells are generally small and contain numerous small vacuoles. As water accumulates in these vacuoles, they tend to coalesce, forming a large central vacuole. A plant cell increases in size mainly by adding water to this central vacuole. As much as 80% of the volume of a plant cell may be occupied by a large central vacuole containing water, stored food, salts, pigments, and metabolic wastes (see Figs. 4-7 and 4-9). The vacuole

Don Fawcett/Photo Researchers, Inc.

Lysosomes are compartments for digestion

Primary lysosome

Secondary lysosome

5 μm

FIGURE 4-17 Lysosomes The dark vesicles in this TEM are lysosomes, compartments that separate powerful digestive enzymes from the rest of the cell. Primary lysosomes bud off from the Golgi complex. After a lysosome takes in material to be digested, it is known as a secondary lysosome. The large vesicles shown here are secondary lysosomes containing various materials being digested.

may serve as a storage compartment for inorganic compounds. In seeds, vacuoles store molecules such as proteins. Because the vacuole contains a high concentration of solutes (dissolved materials), it takes in water and pushes outward on the cell wall. This hydrostatic pressure, called turgor pressure, provides much of the mechanical strength of plant cells. The vacuole is important in maintaining homeostasis. For example, it helps maintain appropriate pH by taking in excess hydrogen ions. Plants lack organ systems for disposing of toxic metabolic waste products. Plant vacuoles are like lysosomes in that they contain hydrolytic enzymes and break down wastes, as well as unneeded organelles and other cell components. Wastes may be recycled in the vacuole, or they may aggregate and form small crystals inside the vacuole. Compounds that are noxious to herbivores (animals that eat plants) may be stored in some plant vacuoles as a means of defense. Vacuoles are also present in many types of animal cells and in unicellular protists such as protozoa. Most protozoa have food vacuoles, which fuse with lysosomes that digest the food (FIG. 4-18). Some protozoa also have contractile vacuoles, which remove excess water from the cell (discussed in Chapter 26).

Peroxisomes metabolize small organic compounds Peroxisomes are membrane-enclosed organelles containing enzymes that catalyze an assortment of metabolic reactions in

M. I. Walker/Photo Researchers, Inc.

Food vacuoles containing diatoms

15 μm

FIGURE 4-18 LM of food vacuoles This protist, Chilodonella, has ingested many small, photosynthetic protists called diatoms (dark areas) that have been enclosed in food vacuoles. From the number of diatoms scattered about its cell, one might conclude that Chilodonella has a rather voracious appetite.

Mitochondria and chloroplasts are energy-converting organelles

which hydrogen is transferred from various compounds to oxygen (FIG. 4-19). Peroxisomes get their name from the fact that during these oxidation reactions, they produce hydrogen peroxide (H2O2). Hydrogen peroxide detoxifies certain compounds; but if this compound were to escape from the peroxisome, it would damage other membranes in the cell. Peroxisomes contain the enzyme Mitochondria

E. H. Newcomb and S. E. Frederick/Biological Photo Service

Chloroplasts

Peroxisomes 1 μm

catalase, which rapidly splits excess hydrogen peroxide to water and oxygen, rendering it harmless. Peroxisomes are found in large numbers in cells that synthesize, store, or degrade lipids. One of their main functions is to break down fatty acid molecules. Peroxisomes synthesize certain phospholipids that are components of the insulating covering of nerve cells. In fact, certain neurological disorders occur when peroxisomes do not perform this function. Mutations that cause synthesis of abnormal membrane lipids by peroxisomes are linked to some forms of mental retardation. When yeast cells are grown in an alcohol-rich medium, they manufacture large peroxisomes containing an enzyme that degrades the alcohol. Peroxisomes in human liver and kidney cells detoxify certain toxic compounds, including ethanol, the alcohol in alcoholic beverages. In plant seeds, specialized peroxisomes, called glyoxysomes, contain enzymes that convert stored fats to sugars. The sugars are used by the young plant as an energy source and as a component for synthesizing other compounds. Animal cells lack glyoxysomes and cannot convert fatty acids into sugars.

The energy a cell obtains from its environment is usually in the form of chemical energy in food molecules (such as glucose) or in the form of light energy. These types of energy must be converted to forms that cells can use more conveniently. Some energy conversions occur in the cytosol, but other types take place in mitochondria and chloroplasts, organelles specialized to facilitate the conversion of energy from one form to another. Chemical energy is most commonly stored in ATP. Recall from (Chapter 3) that the chemical energy of ATP can be used to drive a variety of chemical reactions in the cell. FIGURE 4-20 summarizes the main activities that take place in mitochondria, found in almost all eukaryotic cells, and in chloroplasts, found only in algae and certain plant cells. Mitochondria and chloroplasts grow and reproduce themselves. They contain small amounts of DNA that code for a small number of the proteins found in these organelles. These proteins are synthesized by mitochondrial or chloroplast ribosomes. Interestingly, these ribosomes are similar to the ribosomes of prokaryotes. The existence of a separate set of ribosomes and DNA molecules in mitochondria and chloroplasts and their similarity in size to many bacteria provide support for serial endosymbiosis (discussed in Chapters 21 and 26; see Figs. 21-8 and 26-2). According to this hypothesis, mitochondria and chloroplasts evolved from prokaryotes that took up residence inside larger eukaryotic cells. Eventually, these symbiotic prokaryotes lost the ability to function as autonomous organisms.

FIGURE 4-19 Peroxisomes

Mitochondria make ATP through cellular respiration

In this TEM of a tobacco (Nicotiana tabacum) leaf cell, peroxisomes are in close association with chloroplasts and mitochondria. These organelles may cooperate in carrying out some metabolic processes.

Virtually all eukaryotic cells (plant, animal, fungal, and protist) contain complex organelles called mitochondria (sing., mito-

KEY POINT

Mitochondria and chloroplasts convert energy into forms that can be used by cells.

Aerobic respiration Mitochondria (most eukaryotic cells)

Photosynthesis Chloroplasts (some plant and algal cells) Light

Glucose + O2

ATP

+

CO2

CO2

H2O

H2O

+

ATP

O2 + Glucose

FIGURE 4-20 Cellular respiration and photosynthesis

chondrion). These organelles are the site of aerobic respiration, an oxygen-requiring process that includes most of the reactions that convert the chemical energy present in certain foods to ATP (discussed in Chapter 8). During aerobic respiration, carbon and oxygen atoms are removed from food molecules, such as glucose, and converted to carbon dioxide and water. Mitochondria are most numerous in cells that are very active and therefore have high energy requirements. More than 1000 mitochondria have been counted in a single liver cell! These organelles vary in size, ranging from 2 to 8 μm in length, and change size and shape rapidly. Mitochondria usually give rise to other mitochondria by growth and subsequent division. Each mitochondrion is enclosed by a double membrane, which forms two different compartments within the organelle: the intermembrane space and the matrix (FIG. 4-21; we will provide more detailed descriptions of mitochondrial structure in Chapter 8). The intermembrane space is the compartment formed between the outer and inner mitochondrial membranes. The matrix, the compartment enclosed by the inner mitochondrial membrane, contains enzymes that break down food molecules and convert their energy to other forms of chemical energy. The outer mitochondrial membrane is smooth and allows many small molecules to pass through it. By contrast, the inner mitochondrial membrane has numerous folds and strictly regulates the types of molecules that can move across it. The folds, called cristae (sing., crista), extend into the matrix. Cristae greatly increase the surface area of the inner mitochondrial membrane, providing a surface for the chemical reactions that transform the chemical energy in food molecules into the energy of ATP. The membrane contains the enzymes and other proteins needed for these reactions. Mitochondria play an important role in programmed cell death, or apoptosis. Unlike necrosis, which is uncontrolled cell

in some plant and algal cells, converts light energy to ATP and to other forms of chemical energy. This energy is used to synthesize glucose from carbon dioxide and water.

Outer mitochondrial membrane

Inner mitochondrial membrane

Bloom and Fawcett Textbook of Histology

Cellular respiration takes place in the mitochondria of virtually all eukaryotic cells. In this process, some of the chemical energy in glucose is transferred to ATP. Photosynthesis, which is carried out in chloroplasts

Matrix Cristae

0.25 μm

FIGURE 4-21 Animated Mitochondria Aerobic respiration takes place within mitochondria. Cristae are evident in the TEM as well as in the drawing. The drawing shows the relationship between the inner and outer mitochondrial membranes.

death that causes inflammation and damages other cells, apoptosis is a normal part of development and maintenance. For example, during the metamorphosis of a tadpole to a frog, the cells of the tadpole tail must die. The hand of a human embryo is webbed until apoptosis destroys the tissue between the fingers. Cell death also occurs in the adult. For example, cells that are no longer functional because they have aged or become damaged are destroyed and replaced by new cells. Mitochondria initiate cell death in several different ways. For example, they can interfere with energy metabolism or activate enzymes that mediate cell destruction. When a mitochondrion is injured, large pores open in its membrane, and cytochrome c, a protein important in energy conversions, is released into the cytoplasm. Cytochrome c triggers apoptosis by activating enzymes known as caspases, which cut up vital compounds in the cell. Inappropriate inhibition of apoptosis may contribute to a variety of diseases, including cancer. On the other hand, too much apoptosis may deplete needed cells and lead to death of brain cells associated with Alzheimer’s disease, Parkinson’s disease, and stroke. Mutations that promote apoptosis may be an important mechanism in mammalian aging. Pharmaceutical companies are developing drugs that block apoptosis. However, cell dynamics are extremely complex, and blocking apoptosis could lead to a worse fate, including necrosis. Each mitochondrion in a mammalian cell has 5 to 10 identical, circular molecules of DNA, accounting for up to 1% of the total DNA in the cell. Mitochondrial DNA mutates far more frequently than nuclear DNA. Mutations in mitochondrial DNA have been associated with certain genetic diseases, including a form of young adult blindness, and certain types of progressive muscle degeneration. Mitochondria also affect health and aging by leaking electrons. Inner membrane

Chloroplasts convert light energy to chemical energy through photosynthesis Certain plant and algal cells carry out photosynthesis, a set of reactions during which light energy is transformed into the chemical energy of glucose and other carbohydrates. Chloroplasts are organelles that contain chlorophyll, a green pigment that traps light energy for photosynthesis. Chloroplasts also contain a variety of light-absorbing yellow and orange pigments known as carotenoids (see Chapter 3). A unicellular alga may have only a single large chloroplast, whereas a leaf cell may have 20 to 100. Chloroplasts tend to be somewhat larger than mitochondria, with lengths usually ranging from about 5 to 10 μm or longer. Chloroplasts are typically disc-shaped structures and, like mitochondria, have a system of folded membranes (FIG. 4-22; we will provide a more detailed description of chloroplast structure in Chapter 9). Two membranes enclose the chloroplast and separate it from the cytosol. The inner membrane encloses a fluid-filled space called the stroma, which contains enzymes. These enzymes produce carbohydrates from carbon dioxide and water, using energy trapped from sunlight. A system of internal membranes, which consists of an interconnected set of flat, disclike sacs called thylakoids, is suspended in the stroma. The thylakoids are arranged in stacks called grana (sing., granum). The thylakoid membrane encloses the innermost compartments within the chloroplast, the thylakoid lumen. Chlorophyll is present in the thylakoid membrane, which like the inner mitoStroma

E. H. Newcomb & W. P. Wergin, Biological Photo Service

Outer membrane

These electrons form free radicals, which are toxic, highly reactive compounds with unpaired electrons. The electrons bond with other compounds in the cell, interfering with normal function.

1 μm

Intermembrane space

Thylakoid membrane

Thylakoid lumen

Granum (stack of thylakoids)

FIGURE 4-22 Animated A chloroplast, the organelle of photosynthesis The TEM shows part of a chloroplast from a corn leaf cell. Chlorophyll and other photosynthetic pigments are in the thylakoid membranes. One granum is cut open to show the thylakoid lumen. The inner chloroplast membrane may or may not be continuous with the thylakoid membrane (as shown).

KEY POINT

The cytoskeleton consists of networks of several types of fibers that support the cell and are important in cell movement

Plasma membrane Microfilament Intermediate filament Microtubule

© Dr. Torsten Wittmann/Photo Researchers, Inc.

chondrial membranes, is involved in the formation of ATP. Energy absorbed from sunlight by the chlorophyll molecules excites electrons; the energy in these excited electrons is then used to produce ATP and other molecules that transfer chemical energy. Chloroplasts belong to a group of organelles, known as plastids, that produce and store food materials in cells of plants and algae. All plastids develop from proplastids, precursor organelles found in less specialized plant cells, particularly in growing, undeveloped tissues. Depending on the specific functions a cell will eventually have, its proplastids can develop into a variety of specialized mature plastids. These are extremely versatile organelles; in fact, under certain conditions even mature plastids can convert from one form to another. Chloroplasts are produced when proplastids are stimulated by exposure to light. Chromoplasts contain pigments that give certain flowers and fruits their characteristic colors; these colors attract animals that serve as pollinators or as seed dispersers. Leukoplasts are unpigmented plastids; they include amyloplasts (see Fig. 3-9), which store starch in the cells of many seeds, roots, and tubers (such as white potatoes).

Review ■

■ ■

■ ■

How do the structure and function of rough ER differ from those of smooth ER? What are the functions of the Golgi complex? What sequence of events must take place for a protein to be manufactured and then secreted from the cell? How are chloroplasts like mitochondria? How are they different? Draw and label a chloroplast and a mitochondrion.

4.6 THE CYTOSKELETON ■ ■ LEARNING OBJECTIVES

FIGURE 4-23 Animated The cytoskeleton The cytoskeleton of eukaryotic cells consists of networks of several types of fibers, including microtubules, microfilaments, and intermediate filaments. The cytoskeleton contributes to the shape of the cell, anchors organelles, and sometimes rapidly changes shape during cell locomotion. This fluorescent LM shows the cytoskeleton of two fibroblast cells (microtubules, yellow; microfilaments, blue; nuclei, green).

12 Describe the structure and functions of the cytoskeleton. 13 Compare cilia and flagella, and describe their functions.

Scientists watching cells growing in the laboratory see that they frequently change shape and that many types of cells move about. The cytoskeleton, a dense network of protein fibers, gives cells mechanical strength, shape, and their ability to move (FIG. 4-23). The cytoskeleton also functions in cell division and in the transport of materials within the cell. The cytoskeleton is highly dynamic and constantly changing. Its framework is made of three types of protein filaments: microtubules, microfilaments, and intermediate filaments. Both microfilaments and microtubules are formed from beadlike, globular protein subunits, which can be rapidly assembled and disassembled. Intermediate filaments are made from fibrous protein subunits and are more stable than microtubules and microfilaments.

Microtubules are hollow cylinders Microtubules, the thickest filaments of the cytoskeleton, are rigid, hollow rods about 25 nm in outside diameter and up to several micrometers in length. In addition to playing a structural role in the

cytoskeleton, these extremely adaptable structures are involved in the movement of chromosomes during cell division. They serve as tracks for several other kinds of intracellular movement and are the major structural components of cilia and flagella—specialized structures used in some cell movements. Microtubules consist of two forms of the protein tubulin: α-tubulin and β-tubulin. These proteins combine to form a dimer. (Recall from Chapter 3 that a dimer forms from the association of two simpler units, referred to as monomers.) A microtubule elongates by the addition of tubulin dimers (FIG. 4-24). Microtubules are disassembled by the removal of dimers, which are recycled to form new microtubules. Each microtubule has polarity, and its two ends are referred to as plus and minus. The plus end elongates more rapidly. Other proteins are important in microtubule function. Microtubule-associated proteins (MAPs) are classified into two groups: structural MAPs and motor MAPs. Structural MAPs may help regulate microtubule assembly, and they cross-link microtubules to other cytoskeletal polymers. Motor MAPs use ATP energy to produce movement.

Dimer on

secretory vesicles, attach to microtubules. The microtubules serve as tracks along which organelles move to different cell locations. One motor protein, kinesin, moves organelles toward the plus end of a microtubule (FIG. 4-25). Dynein, another motor protein, transports organelles in the opposite direction, toward the minus end. This dynein movement is referred to as retrograde transport. A protein complex called dynactin is also required for retrograde transport. Dynactin is an adapter protein that links dynein to the microtubule and the organelle. At times, for example, during peroxisome transport, kinesins and dyneins may work together.

α-Tubulin Plus end β-Tubulin

Centrosomes and centrioles function in cell division Minus end

Dimers off

Nancy Kedersha

(a) Microtubules are manufactured in the cell by adding dimers of α-tubulin and β-tubulin to an end of the hollow cylinder. Notice that the cylinder has polarity. The end shown at the top of the figure is the fast-growing, or plus, end; the opposite end is the minus end. Each turn of the spiral requires 13 dimers.

50 μm

(b) Fluorescent LM showing microtubules in green. A microtubule-organizing center (pink dot) is visible beside or over most of the cell nuclei (blue).

FIGURE 4-24 Organization of microtubules

What are the mechanisms by which organelles and other materials move within the cell? Nerve cells typically have long extensions called axons that transmit signals to other nerve cells, muscle cells, or cells that produce hormones. Because of the axon’s length and accessibility and because other cells use similar transport mechanisms, researchers have used the axon as a model for studying the transport of organelles within the cell. They have found that many organelles, including mitochondria and transport and

For microtubules to act as a structural framework or participate in cell movement, they must be anchored to other parts of the cell. In nondividing cells, the minus ends of microtubules appear to be anchored in regions called microtubule-organizing centers (MTOCs). In animal cells, the main MTOC is the cell center, or centrosome, a structure that is important in cell division. In many cells, including almost all animal cells, the centrosome contains two structures called centrioles (FIG. 4-26). These structures are oriented within the centrosome at right angles to each other. They are known as 9 × 3 structures because they consist of nine sets of three attached microtubules arranged to form a hollow cylinder. The centrioles are duplicated before cell division and may play a role in some types of microtubule assembly. Most plant cells and fungal cells have an MTOC but lack centrioles. This suggests that centrioles are not essential to most microtubule assembly processes and that alternative assembly mechanisms are present. The ability of microtubules to assemble and disassemble rapidly is seen during cell division, when much of the cytoskeleton disassembles (discussed in Chapter 10). At that time tubulin subunits organize into a structure called the mitotic spindle, which serves as a framework for the orderly distribution of chromosomes during cell division.

Cilia and flagella are composed of microtubules Thin, movable structures, important in cell movement, project from surfaces of many cells. If a cell has one, or only a few, of these appendages and if they are long (typically about 200 μm) relative to the size of the cell, they are called flagella. If the cell has many short (typically 2 to 10 μm long) appendages, they are called cilia (sing., cilium). Cilia and flagella help unicellular and small multicellular organisms move through a watery environment. In animals and certain plants, flagella serve as the tails of sperm cells. In animals, cilia occur on the surfaces of cells that line internal ducts of the body (such as respiratory passageways). Cells use cilia to move liquids and particles across the cell surface. Investigators have shown that cilia also serve as the cell’s antenna and play important roles in cell signaling. Eukaryotic cilia and flagella are structurally alike (but different from bacterial flagella). Each consists of a slender, cylindrical stalk covered by an extension of the plasma membrane. The core

Vesicle Kinesin receptor Kinesin ATP

ATP MTOC

Minus end

Plus end Centrioles

Microtubule does not move

FIGURE 4-25 Animated A model of a kinesin motor

of the stalk contains a group of microtubules arranged so there are nine attached pairs of microtubules around the circumference and two unpaired microtubules in the center (FIG. 4-27). This 9 + 2 arrangement of microtubules is characteristic of virtually all eukaryotic cilia and flagella. The microtubules in cilia and flagella move by sliding in pairs past each other. The sliding force is generated by dynein proteins, powered by ATP. The dynein “feet” move the microtubule pairs by forming and breaking cross bridges on adjacent pairs of microtubules. Each microtubule pair “walks” along its neighbor. Flexible linking proteins between microtubule pairs prevent microtubules from sliding very far. As a result, the motor action causes the microtubules to bend, resulting in a beating motion (see Fig. 4-27e). Cilia typically move like the arms of a swimmer, alternating a power stroke in one direction with a recovery stroke in the opposite direction. They exert a force that is parallel to the cell surface. In contrast, a flagellum moves like a whip, exerting a force perpendicular to the cell surface. Each cilium or flagellum is anchored in the cell by a basal body, which has nine sets of three attached microtubules in a cylindrical array (9 × 3 structure). The basal body appears to be the organizing structure for the cilium or flagellum when it first begins to form. However, experiments have shown that as growth proceeds, the tubulin subunits are added much faster to the tips of the microtubules than to the base. Basal bodies and centrioles may be functionally related as well as structurally similar. In fact, centrioles are typically found in the cells of eukaryotic organisms that produce flagellated or ciliated cells; these include animals, certain protists, a few fungi, and a few plants. Both basal bodies and centrioles replicate themselves. Almost every vertebrate cell has a primary cilium, a single cilium on the cell surface that serves as a cellular antenna. The primary cilium has receptors on its surface that bind with specific molecules outside the cell or on surfaces of other cells. Recent research indicates that primary cilia play important roles in many signaling pathways that regulate growth and specialization of cells during embryonic development. Primary cilia also help maintain healthy tissues. Malfunction of primary cilia has been associated

B. F. King/Biological Photo Service

A kinesin molecule attaches to a specific receptor on the vesicle. Energy from ATP powers the kinesin molecule so that it changes its conformation and “walks” along the microtubule, carrying the vesicle along. (Size relationships are exaggerated for clarity.)

0.25 μm

(a) In the TEM, the centrioles are positioned at right angles to each other, near the nucleus of a nondividing animal cell.

(b) Note the 9 × 3 arrangement of microtubules. The centriole on the right has been cut transversely.

FIGURE 4-26 Centrioles

with several human disorders, including developmental defects, degeneration of the retina, and polycystic kidney disease.

Microfilaments consist of intertwined strings of actin Microfilaments, also called actin filaments, are flexible, solid fibers about 7 nm in diameter. Each microfilament consists of two intertwined polymer chains of beadlike actin molecules (FIG. 4-28). Microfilaments are linked with one another and with other proteins by linker proteins. They form bundles of fibers that provide mechanical support for various cell structures. In many cells, a network of microfilaments is visible just inside the plasma membrane, a region called the cell cortex. Microfilaments give the cell cortex a gel-like consistency compared to the

KEY POINT

A cilium consists of a 9 + 2 arrangement of microtubules surrounded by the plasma membrane; dynein proteins move the microtubules by forming and breaking cross bridges on adjacent pairs of microtubules. Outer pair of microtubules Dynein Dennis Kunkel

Plasma membrane Central microtubules

0.5 μm

(d) This 3-D representation shows nine attached microtubule pairs (doublets) arranged in a cylinder, with two unattached microtubules in the center. The dynein “feet,” shown widely spaced for clarity, are actually much closer together along the longitudinal axis.

Dr. David M. Phillips/ Visuals Unlimited

Dennis Kunkel

(b) TEM of cross sections through cilia showing 9 + 2 arrangement of microtubules.

0.5 μm

(a) TEM of a longitudinal section through cilia and basal bodies of the freshwater protist Paramecium. Some of the interior microtubules are visible.

+

+

+

+

+

+

+

+

ATP

0.5 μm Microtubular bend

(c) TEM of cross section through basal body showing 9 × 3 structure.

Linking proteins

(e) The dynein “feet” move the microtubules so that one microtubule pair slides along an adjacent pair. Flexible linking proteins between microtubule pairs prevent microtubules from sliding very far. Instead, the motor action causes the microtubules to bend, resulting in a beating motion.

Dynein Pair of microtubules

–

–

–

–

–

–

–

–

FIGURE 4-27 Animated Structure and movement of cilia

more fluid state of the cytosol deeper inside the cell. The microfilaments in the cell cortex help determine the shape of the cell and are important in its movement. Microfilaments themselves cannot contract, but they can generate movement by rapidly assembling and disassembling. Muscle cells have two types of specialized filaments, one composed mainly of the protein myosin, and another composed mainly of the protein actin. ATP bound to myosin provides energy for muscle contraction. When ATP is hydrolyzed to ADP, myosin binds to actin and causes the microfilament to slide. When thousands of filaments slide in this way, the muscle cell shortens. Thus, ATP, actin, and myosin generate the forces that contract muscles (discussed in Chapter 40). In nonmuscle cells, actin also associates with myosin, forming contractile structures involved in various cell movements. For example, in animal cell division, contraction of a ring of actin asso-

ciated with myosin constricts the cell, forming two daughter cells (discussed in Chapter 10). Some cells change their shape quickly in response to changes in the outside environment. Amoebas, human white blood cells, and cancer cells are among the many cell types that can creep along a surface, a process that includes changes in shape. Such responses depend on external signals that affect microfilament, as well as microtubule, assembly. Actin filaments push the plasma membrane outward, forming cytoplasm-filled bulges called pseudopodia (“false feet”). The pseudopodia adhere to the surface. Contractions of microfilaments at the opposite end of the cell force the cytoplasm forward in the direction of locomotion. Microtubules, myosin, and other proteins also appear necessary for cell creeping. As mentioned earlier in the chapter, some types of cells have microvilli, projections of the plasma membrane that increase the surface area of the cell for transporting materials across the plasma

Protofilament 7 nm

Protein subunits

(a) A microfilament consists of two intertwined strings of beadlike actin molecules.

Intermediate filament

K. G. Murti/Visuals Unlimited

Nancy Kedersha

(a) Intermediate filaments are flexible rods about 10 nm in diameter. Each intermediate filament consists of components, called protofilaments, composed of coiled protein subunits.

100 μm

(b) Many bundles of microfilaments (green) are evident in this fluorescent LM of fibroblasts, cells found in connective tissue. 100 μm

FIGURE 4-28 Microfilaments

(b) Intermediate filaments are stained green in this human cell isolated from a tissue culture.

membrane. Composed of bundles of microfilaments, microvilli extend and retract as the microfilaments assemble and disassemble.

Intermediate filaments help stabilize cell shape Intermediate filaments are tough, flexible fibers about 10 nm in diameter (FIG. 4-29). They provide mechanical strength and help stabilize cell shape. These filaments are abundant in regions of a cell that may be subject to mechanical stress applied from outside the cell. Intermediate filaments prevent the cell from stretching excessively in response to outside forces. Certain proteins cross-link intermediate filaments with other types of filaments and mediate interactions between them. All eukaryotic cells have microtubules and microfilaments, but only some animal groups, including vertebrates, are known to have intermediate filaments. Even when present, intermediate filaments vary widely in protein composition and size among different cell types and different organisms. Examples of intermediate filaments are the keratins found in the epithelial cells of vertebrate skin and neurofilaments found in vertebrate nerve cells. Certain mutations in genes coding for intermediate filaments weaken the cell and are associated with several diseases. For example, in the neurodegenerative disease amyotrophic lateral sclerosis (ALS, or Lou Gehrig’s disease), abnormal neurofilaments have been identified in nerve cells that control muscles. This condition

FIGURE 4-29 Intermediate filaments

interferes with normal transport of materials in the nerve cells and leads to degeneration of the cells. The resulting loss of muscle function is typically fatal.

Review ■ ■

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What are the main functions of the cytoskeleton? How are microfilaments and microtubules similar? How are they different? How are cilia and flagella similar? How are they different?

4.7 CELL COVERINGS ■ ■ LEARNING OBJECTIVE 14 Describe the glycocalyx, extracellular matrix, and cell wall.

Many cells are surrounded by a glycocalyx, or cell coat, formed by polysaccharide side chains of proteins and lipids that are part of the plasma membrane. The glycocalyx protects the cell and may help keep other cells at a distance. Certain molecules of the glycocalyx

Biophoto Associates

enable cells to recognize one another, to make contact, and in some cases to form adhesive or communicating associations (discussed in Chapter 5). Other molecules of the cell coat contribute to the mechanical strength of Collagen multicellular tissues. Many animal cells are also surrounded by an extraFibronectins cellular matrix (ECM), which they secrete. The ECM Extracellular consists of a gel of carbohydrates and fibrous proteins matrix (FIG. 4-30). The main structural protein in the ECM is Integrin collagen, a protein that forms very tough fibers (see Figure 3-22b). Certain glycoproteins of the ECM, called Intermediate filament fibronectins, organize the matrix and help cells attach Plasma to it. Fibronectins bind to protein receptors that extend membrane Microtubules from the plasma membrane. Integrins are receptor proteins in the plasma memCytosol brane. They maintain adhesion between the ECM and the intermediate filaments and microfilaments inside the cell. These proteins activate many cell signaling pathways FIGURE 4-30 The extracellular matrix (ECM) that communicate information from the ECM, and they Fibronectins, glycoproteins of the ECM, bind to integrins and other receptors in the control signals inside the cell that regulate the differentiaplasma membrane. tion and survival of the cell. When cells are not appropriately anchored, apoptosis is initiated. Integrins are also important in organizing the cytoskeleton so that cells assume a definite shape. The cells of most bacteria, archaea, fungi, and plant cells are surrounded by a cell wall. Plant cells have thick cell walls that conCell 1 tain tiny fibers composed of the polysaccharide cellulose (see Fig. 3-10). Other polysaccharides in the plant cell wall form cross links between the bundles of cellulose fibers. Cell walls provide Middle structural support, protect plant cells from disease-causing organlamella isms, and keep excess water out of cells so they do not burst. A growing plant cell secretes a thin, flexible primary cell wall. Primary cell wall As the cell grows, the primary cell wall increases in size (FIG. 4-31). After the cell stops growing, either new wall material is secreted that Multiple layers of thickens and solidifies the primary wall, or multiple layers of a secsecondary cell wall ondary cell wall with a different chemical composition are formed between the primary wall and the plasma membrane. Wood is Cell 2 made mainly of secondary cell walls. The middle lamella, a layer of gluelike polysaccharides called pectins, lies between the primary cell walls of adjacent cells. The middle lamella causes the cells to adhere tightly to one another. (For more information on plant cell walls, 2.5 μm see the discussion of the ground tissue system in Chapter 33.)

Review What are the functions of the glycocalyx? How do the functions of fibronectins and integrins differ? What is the main component of plant cell walls? How are plant cell walls formed?

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FIGURE 4-31 Animated Plant cell walls The cell walls of two adjacent plant cells are labeled in this TEM. The cells are cemented together by the middle lamella, a layer of gluelike polysaccharides called pectins. A growing plant cell first secretes a thin primary wall that is flexible and can stretch as the cell grows. The thicker layers of the secondary wall are secreted inside the primary wall after the cell stops elongating.

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

4.1 (page 75) 1 Describe the cell theory and relate it to the evolution of life. ■

The cell theory holds that (1) cells are the basic living units of organization and function in all organisms and (2) all cells come from

other cells. It explains that the ancestry of all the cells alive today can be traced back to ancient times. Evidence that all living cells have evolved from a common ancestor is supported by the basic similarities in their structures and in their molecular composition.

Test yourself on the structure of prokaryotic and eukaryotic cells by clicking on the interaction in CengageNOW.

2 Summarize the relationship between cell organization and homeostasis. ■ The organization of cells is important in maintaining homeostasis, an appropriate internal environment. ■ Every cell is surrounded by a plasma membrane that separates it from its external environment. The plasma membrane allows the cell to maintain internal conditions that may be very different from those of the outer environment. The plasma membrane also allows the cell to exchange materials with its outer environment. Cells have many organelles, internal structures that carry out specific functions that help maintain homeostasis. 3 Explain the relationship between cell size and homeostasis. ■ A critical factor in determining cell size is the ratio of the plasma membrane (surface area) to the cell’s volume. The plasma membrane must be large enough relative to the cell volume to regulate the passage of materials into and out of the cell. For this reason, most cells are microscopic. ■ Cell size and shape are related to function and are limited by the need to maintain homeostasis.

6 Describe three functions of cell membranes. ■

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Learn more about the endomembrane system by clicking on the figure in CengageNOW.

4.4 (page 86) 7 Describe the structure and functions of the nucleus. ■

4.2 (page 77) 4 Describe methods that biologists use to study cells, including microscopy and cell fractionation. Biologists use light microscopes, electron microscopes, and a variety of chemical methods to study cells and learn about cell structure. The electron microscope has superior resolving power, enabling investigators to see details of cell structures not observable with conventional microscopes. ■ Cell biologists use the technique of cell fractionation for purifying organelles to gain information about the function of cell structures.

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4.3 (page 81) 5 Compare and contrast the general characteristics of prokaryotic and eukaryotic cells, and contrast plant and animal cells. Prokaryotic cells are enclosed by a plasma membrane but have little or no internal membrane organization. They have a nuclear area rather than a membrane-enclosed nucleus. Prokaryotic cells typically have a cell wall and ribosomes and may have propellerlike flagella. ■ Eukaryotic cells have a membrane-enclosed nucleus, and their cytoplasm contains a variety of organelles; the fluid component of the cytoplasm is the cytosol. Plant cells differ from animal cells in that plant cells have rigid cell ■ walls, plastids, and large vacuoles, which are important in plant growth and development. Cells of most plants lack centrioles. ■

Membranes divide the eukaryotic cell into compartments, allowing it to conduct specialized activities within small areas of the cytoplasm, concentrate reactants, and organize metabolic reactions. Small membrane-enclosed sacs, called vesicles, transport materials between compartments. Membranes are important in energy storage and conversion. A system of interacting membranes forms the endomembrane system.

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The nucleus, the control center of the eukaryotic cell, contains genetic information coded in DNA. The nucleus is bounded by a nuclear envelope, consisting of a double membrane perforated with nuclear pores that communicate with the cytoplasm. DNA in the nucleus associates with protein to form chromatin, which is organized into chromosomes. During cell division, the chromosomes condense and become visible as threadlike structures. The nucleolus is a region in the nucleus that is the site of ribosomal RNA synthesis and ribosome assembly.

4.5 (page 89) 8 Distinguish between smooth and rough endoplasmic reticulum in terms of both structure and function. ■ The endoplasmic reticulum (ER) is a network of folded internal membranes in the cytosol. Smooth ER is the site of lipid synthesis, calcium ion storage, and detoxifying enzymes. ■ Rough ER is studded along its outer surface with ribosomes that manufacture polypeptides. Polypeptides synthesized on rough ER may be moved into the ER lumen, where they are assembled into proteins and modified by the addition of a carbohydrate or lipid. These proteins may then be transferred to other compartments within the cell by small transport vesicles that bud off from the ER membrane. 9 Trace the path of proteins synthesized in the rough endoplasmic reticulum as they are processed, modified, and sorted by the Golgi complex and then transported to specific destinations. ■ The Golgi complex consists of stacks of flattened membranous sacs called cisternae that process, sort, and modify proteins synthesized on the ER. The Golgi complex also manufactures lysosomes. ■ Glycoproteins are transported from the ER to the cis face of the Golgi complex by transport vesicles, which are formed by membrane budding. The Golgi complex modifies carbohydrates and lipids that were added to proteins by the ER and packages them in vesicles. ■ Glycoproteins exit the Golgi at its trans face. The Golgi routes some proteins to the plasma membrane for export from the cell. Others are transported to lysosomes or other organelles within the cytoplasm. 10 Describe the functions of lysosomes, vacuoles, and peroxisomes. ■ Lysosomes contain enzymes that break down worn-out cell structures, bacteria, and debris taken into cells. Vacuoles store materials, water, and wastes. They maintain hydro■ static pressure in plant cells. ■ Peroxisomes are important in lipid metabolism and detoxify harmful compounds such as ethanol. They produce hydrogen peroxide but contain the enzyme catalase, which degrades this toxic compound.

11 Compare the functions of mitochondria and chloroplasts, and discuss

4.6 (page 97)

ATP synthesis by each of these organelles. ■ Mitochondria, organelles enclosed by a double membrane, are the sites of aerobic respiration. The inner membrane is folded, forming cristae that increase its surface area. ■ The cristae and the compartment enclosed by the inner membrane, the matrix, contain enzymes for the reactions of aerobic respiration. During aerobic respiration, nutrients are broken down in the presence of oxygen. Energy captured from nutrients is packaged in ATP, and carbon dioxide and water are produced as byproducts. Outer mitochondrial membrane

Inner mitochondrial membrane

12 Describe the structure and functions of the cytoskeleton. ■

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Bloom and Fawcett Textbook of Histology

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Matrix Cristae

0.25 μm

Learn more about the cytoskeleton by clicking on the figure in CengageNOW.

13 Compare cilia and flagella, and describe their functions. ■

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Plastids are organelles that produce and store food in the cells of plants and algae. Chloroplasts are plastids that carry out photosynthesis. The inner membrane of the chloroplast encloses a fluid-filled space, the stroma. Grana, stacks of interconnected disclike membranous sacs called thylakoids, are suspended in the stroma. During photosynthesis, chlorophyll, the green pigment found in the thylakoid membranes, traps light energy. This energy is converted to chemical energy in ATP and used to synthesize carbohydrates from carbon dioxide and water.

Cilia and flagella are thin, movable structures that project from the cell surface and function in movement. Each consists of a 9 + 2 arrangement of microtubules, and each is anchored in the cell by a basal body that has a 9 × 3 organization of microtubules. Cilia are short, and flagella are long.

4.7 (page 101) 14 Describe the glycocalyx, extracellular matrix, and cell wall. ■

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The cytoskeleton is a dynamic internal framework made of microtubules, microfilaments, and intermediate filaments. The cytoskeleton provides structural support and functions in various types of cell movement, including transport of materials in the cell. Microtubules are hollow cylinders assembled from subunits of the protein tubulin. In cells that are not dividing, the minus ends of microtubules are anchored in microtubule-organizing centers (MTOCs). The main MTOC of animal cells is the centrosome, which usually contains two centrioles. Each centriole has a 9 × 3 arrangement of microtubules. Microfilaments, or actin filaments, formed from subunits of the protein actin, are important in cell movement. Intermediate filaments strengthen the cytoskeleton and stabilize cell shape.

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Most cells are surrounded by a glycocalyx, or cell coat, formed by polysaccharides extending from the plasma membrane. Many animal cells are also surrounded by an extracellular matrix (ECM) consisting of carbohydrates and protein. Fibronectins are glycoproteins of the ECM that bind to integrins, receptor proteins in the plasma membrane. Cells of most bacteria, archaea, fungi, and plant cells are surrounded by a cell wall made mainly of carbohydrates. Plant cells secrete cellulose and other polysaccharides that form rigid cell walls.

T E S T YO U R U N D E R S TA N D I N G 1. The ability of a microscope to reveal fine detail is known as (a) magnification (b) resolving power (c) cell fractionation (d) transmission microscopy (e) phase contrast

5. Which of the following is/are most closely associated with protein synthesis? (a) ribosomes (b) smooth ER (c) mitochondria (d) microfilaments (e) lysosomes

2. A plasma membrane is characteristic of (a) all cells (b) prokaryotic cells only (c) eukaryotic cells only (d) animal cells only (e) eukaryotic cells except plant cells

6. Which of the following is/are most closely associated with the breakdown of ingested material? (a) ribosomes (b) smooth ER (c) mitochondria (d) microfilaments (e) lysosomes

3. Detailed information about the shape and external features of a specimen can best be obtained by using a (a) differential centrifuge (b) fluorescence microscope (c) transmission electron microscope (d) scanning electron microscope (e) light microscope

7. Which of the following are most closely associated with photosynthesis? (a) basal bodies (b) smooth ER (c) cristae (d) thylakoids (e) MTOCs

4. Which of the following structures would not be found in prokaryotic cells? (a) cell wall (b) ribosomes (c) nuclear area (d) nucleus (e) propeller-like flagellum

8. Use the choices of numbered sequences to select the sequence that most accurately describes information flow in the eukaryotic cell. 1. DNA in nucleus 2. RNA 3. mitochondria 4. protein synthesis 5. ribosomes (a) 1, 2, 5, 4 (b) 3, 2, 5, 1 (c) 5, 2, 3, 1 (d) 4, 3, 2, 1 (e) 1, 2, 3, 4

9. Use the choices of numbered sequences to select the sequence that most accurately describes glycoprotein processing in the eukaryotic cell. 1. ER 2. ribosomes 3. cis face of Golgi 4. trans face of Golgi 5. transport vesicle 6. grana (a) 1, 6, 2, 3, 4 (b) 2, 1, 3, 4, 5 (c) 1, 2, 4, 3, 5 (d) 2, 1, 4, 3, 5 (e) 2, 1, 4, 5, 1 10. Which of the following organelles contain small amounts of DNA and convert energy? (a) microfilaments and microtubules

(b) lysosomes and peroxisomes (c) ER and ribosomes (d) nucleus and ribosomes (e) mitochondria and chloroplasts 11. Which of the following function(s) in cell movement? (a) microtubules (b) nucleolus (c) grana (d) smooth ER (e) rough ER 12. Label the diagrams of the animal and plant cells. How is the structure of each organelle related to its function? Use Figures 4-9 and 4-10 to check your answers.

CRITICAL THINKING 1. Why does a eukaryotic cell need both membranous organelles and fibrous cytoskeletal components? 2. Describe a specific example of the correlation between cell structure and function. (Hint: Think of mitochondrial structure.) 3. The Acetabularia experiments described in this chapter suggest that DNA is much more stable in the cell than is messenger RNA. Is this advantageous or disadvantageous to the cell? Why? Acetabularia continues to live for a few days after its nucleus is removed. How can you explain this? 4. EVOLUTION LINK. What are the implications of the cell theory for the evolution of organisms?

5. EVOLUTION LINK. Biologists use similarities in cells to trace the evolutionary history of various groups of organisms. Explain their rationale. What do such similarities in cell structure and function tell biologists about the common origin of organisms? Explain. Additional questions are available in CengageNOW at www.cengage.com/ login.

5

Biological Membranes

Nancy Kedersha

Cadherins. The human skin cells shown in this LM were grown in culture and stained with fluorescent antibodies. Cadherins, a group of membrane proteins, are seen as green belts around each cell in this sheet of cells. The nuclei appear as blue spheres; myosin in the cells appears red.

KEY CONCEPTS 5.1 Biological membranes are selectively permeable membranes that help maintain homeostasis in the cell; each cell membrane consists of a fluid bilayer composed of phospholipids in which a variety of proteins are embedded. 5.2 Membrane proteins, and thus membranes, have many

T

he evolution of biological membranes that separate the cell from its external environment was an essential step in the origin of life. Later,

these membranes made the evolution of complex cells possible. The extensive internal membranes of eukaryotic cells form multiple compartments with unique environments that allow highly specialized activities to take place. Because membrane proteins are critical molecules in cell membrane

functions, including transport of materials, enzymatic activity, transfer of information, and recognition of other cells.

activities, they are a major focus of cell membrane research. Some pro-

5.3 Transport proteins move ions and small polar mol-

teins associated with the plasma membrane transport materials, whereas

ecules through cell membranes.

others transmit information or serve as enzymes. Still others, known as

5.4 Many ions and small molecules move through biological membranes by diffusion, the net movement of particles from a region of higher to a region of lower concentrations.

cell adhesion molecules, are important in connecting cells to one another

5.5 Because a cell requires many substances in higher concentrations than their concentrations outside the cell, substances must be actively transported against a concentration gradient; active transport requires a direct expenditure of energy.

to form tissues. The principal cell adhesion molecules in vertebrates and in many invertebrates are cadherins. The genes that code cadherins are found in choanoflagellates, the closest single-celled relatives of animals. These molecules were probably important in the evolution of animals because they provided a means for cells to adhere to one another and to communicate. Cadherins are

5.6 Cells eject products by exocytosis and import materi-

responsible for adhesion between cells that form multicellular sheets.

als by endocytosis; both processes require cells to expend energy.

For example, these membrane proteins form cell junctions important in

5.7 Cells in close contact may form specialized junctions with one another.

maintaining the structure of the epithelium that makes up human skin (see photograph). Cadherins are also important in signaling among cells. An absence of these membrane proteins is associated with the invasiveness of some malignant tumors. In Chapter 4, we discussed a variety of cell organelles and how they interact to perform cell activities. In this chapter, we focus on the structure and functions of the plasma membrane that surrounds the cell, and on the biological membranes that surround many organelles. We first consider what is known about the composition and structure of biological membranes. Then we provide an overview of the many vital functions of cell membranes, including transport of materials and information transfer. We discuss how cells transport various materials, from ions to complex molecules and even bacteria, across membranes. Finally, we examine specialized structures that allow membranes of different cells to interact. Although much of our discussion centers on the structure and functions of plasma membranes, many of the concepts apply to other cell membranes.

5.1 THE STRUCTURE OF BIOLOGICAL MEMBRANES ■ ■ LEARNING OBJECTIVES 1 2 3 4

Evaluate the importance of membranes to the homeostasis of the cell, emphasizing their various functions. Describe the fluid mosaic model of cell membrane structure. Relate properties of the lipid bilayer to properties and functions of cell membranes. Describe the ways that membrane proteins associate with the lipid bilayer.

To carry out the many chemical reactions necessary to sustain life, the cell must maintain an appropriate internal environment. Every cell is surrounded by a plasma membrane that physically separates it from the outside world and defines it as a distinct entity. By regulating passage of materials into and out of the cell, the plasma membrane helps maintain a life-supporting internal environment. As we discussed in Chapter 4, eukaryotic cells are characterized by numerous organelles that are surrounded by membranes. Some of these organelles—including the nuclear envelope, endoplasmic reticulum (ER), Golgi complex, lysosomes, vesicles, and vacuoles—form the endomembrane system, which extends throughout the cell. Biological membranes are complex, dynamic structures made of lipid and protein molecules that are in constant motion. The properties of membranes allow them to perform many vital functions. They regulate the passage of materials, divide the cell into compartments, serve as surfaces for chemical reactions, adhere to and communicate with other cells, and transmit signals between the environment and the interior of the cell. Membranes are also an essential part of energy transfer and storage systems. How do the properties of cell membranes enable the cell to carry on such varied functions?

Long before the development of the electron microscope, scientists knew that membranes consist of both lipids and proteins. Work by researchers in the 1920s and 1930s had provided clues that the core of cell membranes consists of lipids, mostly phospholipids (see Chapter 3).

Phospholipids form bilayers in water Phospholipids are primarily responsible for the physical properties of biological membranes, because certain phospholipids have unique attributes, including features that allow them to form bilayered structures. A phospholipid contains two fatty acid chains linked to two of the three carbons of a glycerol molecule (see Fig. 3-13). The fatty acid chains make up the nonpolar, hydrophobic (“water-fearing”) portion of the phospholipid. Bonded to the third carbon of the glycerol is a negatively charged, hydrophilic (“water-loving”) phosphate group, which in turn is linked to a polar, hydrophilic organic group. Molecules of this type, which have distinct hydrophobic and hydrophilic regions, are called amphipathic molecules. All lipids that make up the core of biological membranes have amphipathic characteristics. Because one end of each phospholipid associates freely with water and the opposite end does not, the most stable orientation for them to assume in water results in the formation of a bilayer structure (FIG. 5-1a). This arrangement allows the hydrophilic heads of the phospholipids to be in contact with the aqueous medium while their oily tails, the hydrophobic fatty acid chains, are buried in the interior of the structure away from the water molecules. Amphipathic properties alone do not predict the ability of lipids to associate as a bilayer. Shape is also important. Phospholipids tend to have uniform widths; their roughly cylindrical shapes, together with their amphipathic properties, are responsible for bilayer formation. In summary, phospholipids form bilayers because the molecules have (1) two distinct regions, one strongly hydrophobic and the other strongly hydrophilic (making them strongly amphipathic) and (2) cylindrical shapes that allow them to associate with water most easily as a bilayer.

Hydrophilic heads

Hydrophobic tails

(a) Phospholipids in water. Phospholipids associate as bilayers in water because they are roughly cylindrical amphipathic molecules. The hydrophobic fatty acid chains are not exposed to water, whereas the hydrophilic phospholipid heads are in contact with water.

(b) Detergent in water. Detergent molecules are roughly cone-shaped amphipathic molecules that associate in water as spherical structures.

FIGURE 5-1 Animated Properties of lipids in water

KEY POINT

The Davson–Danielli model was the accepted view until about 1970 when advances in biology and chemistry led to new findings about biological membranes that were incompatible with this model. The fluid mosaic model is supported by a large body of data. Membrane proteins

Hydrophilic region of protein

Hydrophobic region of protein

Phospholipid bilayer

Phospholipid bilayer

Membrane proteins

(a) The Davson–Danielli model. According to this model, the membrane is a sandwich of phospholipids spread between two layers of protein. Although accepted for many years, this model was shown to be incorrect.

Integral (transmembrane) protein

Peripheral protein

(b) Fluid mosaic model. According to this model, a cell membrane is a fluid lipid bilayer with a constantly changing “mosaic pattern“ of associated proteins.

FIGURE 5-2 Two models of membrane structure

The fluid mosaic model explains membrane structure By examining the plasma membrane of mammalian red blood cells and comparing their surface area with the total number of lipid molecules per cell, early investigators calculated that the membrane is no more than two phospholipid molecules thick. In 1935, these findings, together with other data, led Hugh Davson and James Danielli, working at London’s University College, to propose a model in which they envisioned a membrane as a kind of “sandwich” consisting of a lipid bilayer (a double layer of lipid) between two protein layers (FIG. 5-2a). This useful model had a great influence on the direction of membrane research for more than 20 years. Models are important in the scientific process; good ones not only explain the available data but are testable. Scientists use models to help them develop hypotheses that can be tested experimentally (see Chapter 1). With the development of the electron microscope in the 1950s, cell biologists were able to see the plasma membrane for the first time. One of their most striking observations was how uniform and thin the membranes are. The plasma membrane is no more than 10 nm thick. The electron microscope revealed a three-

layered structure, something like a railroad track, with two dark layers separated by a lighter layer (FIG. 5-3). Their findings seemed to support the protein–lipid–protein sandwich model. During the 1960s, a paradox emerged regarding the arrangement of the proteins. Biologists assumed membrane proteins were uniform and had shapes that would allow them to lie like thin sheets on the membrane surface. But when purified by cell fractionation, the proteins were far from uniform; in fact, they varied widely in composition and size. Some proteins are quite large. How could they fit within a surface layer of a membrane less than 10 nm thick? Cell interior

Plasma membrane

Omikron/Photo Researchers, Inc.

Do you know why detergents remove grease from your hands or from dirty dishes? Many common detergents are amphipathic molecules, each containing a single hydrocarbon chain (like a fatty acid) at one end and a hydrophilic region at the other. These molecules are roughly cone-shaped, with the hydrophilic end forming the broad base and the hydrocarbon tail leading to the point. Because of their shapes, these molecules do not associate as bilayers but instead tend to form spherical structures in water (FIG. 5-1b). Detergents can “solubilize” oil because the oil molecules associate with the hydrophobic interiors of the spheres.

Outside cell

0.1 μm

FIGURE 5-3 TEM of the plasma membrane of a mammalian red blood cell The plasma membrane separates the cytosol (darker region) from the external environment (lighter region). The hydrophilic heads of the phospholipids are the parallel dark lines, and the hydrophobic tails are the light zone between them.

At first, some researchers tried to answer this question by modifyLateral ing the model with the hypothesis that the proteins on the memmovement only brane surfaces were a flattened, extended form, perhaps a β-pleated sheet (see Fig. 3-20b). Other cell biologists found that instead of having sheetlike structures, many membrane proteins are rounded, or globular. Studies of many membrane proteins showed that one region (or Time domain) of the molecule could always be found on one side of the bilayer, whereas another part of the protein might be located on the opposite side. Rather than form a thin surface layer, many membrane proteins extended completely through the lipid bilayer. FIGURE 5-4 Membrane fluidity Thus, the evidence suggested that membranes contain different The ordered arrangement of phospholipid molecules makes the cell membrane a liquid crystal. The hydrocarbon chains are in constant motion, types of proteins of different shapes and sizes that are associated allowing each molecule to move laterally on the same side of the bilayer. with the bilayer in a mosaic pattern. In 1972, S. Jonathan Singer and Garth Nicolson of the Unithem to move along the plane of the membrane (as long as they versity of California at San Diego proposed a model of membrane are not anchored in some way). David Frye and Michael Edidin structure that represented a synthesis of the known properties of elegantly demonstrated this in 1970. They conducted experiments biological membranes. According to their fluid mosaic model, in which they followed the movement of membrane proteins on a cell membrane consists of a fluid bilayer of phospholipid molthe surface of two cells that had been joined (FIG. 5-5). When the ecules in which the proteins are embedded or otherwise associplasma membranes of a mouse cell and a human cell are fused, ated, much like the tiles in a mosaic picture. This mosaic pattern within minutes at least some of the membrane proteins from each is not static, however, because the positions of many of the proteins are constantly changing as they move about like icebergs in a fluid sea of phospholipids. This model has KEY EXPERIMENT provided great impetus to research; QUESTION: Do proteins embedded in a biological membrane move about? it has been repeatedly tested and has been shown to accurately predict HYPOTHESIS: Proteins are able to move laterally in the plasma membrane. the properties of many kinds of cell EXPERIMENT: Larry Frye and Michael Edidin labeled membrane proteins of mouse and human cells membranes. FIGURE 5-2b depicts the using different-colored fluorescent dyes to distinguish between them. Then they fused mouse and huplasma membrane of a eukaryotic cell man cells to produce hybrid cells. according to the fluid mosaic model; Human cell Mouse cell prokaryotic plasma membranes are discussed in Chapter 25.

Biological membranes are two-dimensional fluids An important physical property of phospholipid bilayers is that they behave like liquid crystals. The bilayers are crystal-like in that the lipid molecules form an ordered array with the heads on the outside and fatty acid chains on the inside; they are liquid-like in that despite the orderly arrangement of the molecules, their hydrocarbon chains are in constant motion. Thus, molecules are free to rotate and can move laterally within their single layer (FIG. 5-4). Such movement gives the bilayer the property of a two-dimensional fluid. Under normal conditions a single phospholipid molecule can travel laterally across the surface of a eukaryotic cell in seconds. The fluid qualities of lipid bilayers also allow molecules embedded in

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Labeled membrane proteins. Membrane proteins of mouse cells and human cells were labeled with fluorescent dye markers in two different colors.

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Human-mouse hybrid cell forming. When plasma membranes of mouse cell and human cell were fused, mouse proteins migrated to the human side and human proteins to the mouse side.

3

Proteins randomly distributed. After a short time, mouse and human proteins became randomly distributed through the membrane.

RESULTS AND CONCLUSION: After a brief period of incubation, mouse and human cells intermixed over the surface of the hybrid cells. After about 40 minutes, the proteins of each species had become randomly distributed through the entire hybrid plasma membrane. This experiment demonstrated that proteins in the plasma membrane do move.

FIGURE 5-5 Animated Frye and Edidin’s experiment Source: Frye, L. D., and M. Edidin. “The rapid intermixing of cell surface antigens after formation of mouse-human heterokaryons.” Journal of Cell Science, Vol. 7, 319–335, 1970.

cell migrate and become randomly distributed over the single, continuous plasma membrane that surrounds the joined cells. Frye and Edidin showed that the fluidity of the lipids in the membrane allows many of the proteins to move, producing an ever-changing configuration. Occasionally, with the help of enzymes in the cell membrane, phospholipid molecules flip-flop from one layer to the other. For a membrane to function properly, its lipids must be in a state of optimal fluidity. The membrane’s structure is weakened if its lipids are too fluid. However, many membrane functions, such as the transport of certain substances, are inhibited or cease if the lipid bilayer is too rigid. At normal temperatures, cell membranes are fluid, but at low temperatures the motion of the fatty acid chains is slowed. If the temperature decreases to a critical point, the membrane is converted to a more solid gel state.

KEY POINT

Certain properties of membrane lipids have significant effects on the fluidity of the bilayer. Recall from Chapter 3 that molecules are free to rotate around single carbon-to-carbon covalent bonds. Because most of the bonds in hydrocarbon chains are single bonds, the chains themselves twist more and more rapidly as the temperature rises. The fluid state of the membrane depends on its component lipids. You have probably noticed that when melted butter is left at room temperature it solidifies. Vegetable oils, however, remain liquid at room temperature. Recall from our discussion of lipids in Chapter 3 that animal fats such as butter are high in saturated fatty acids that lack double bonds. In contrast, a vegetable oil may be polyunsaturated, with most of its fatty acid chains having two or more double bonds. At each double bond there is a bend in the molecule that prevents the hydrocarbon chains from coming

According to the fluid mosaic model, a cell membrane is composed of a fluid bilayer of phospholipids in which proteins move about like icebergs in a sea.

Carbohydrate chains

Glycoprotein Carbohydrate chain

Extracellular fluid Hydrophobic

Hydrophilic

Glycolipid

Hydrophobic tails Cholesterol Hydrophilic

α-helix Peripheral protein

Integral proteins

Cytosol

FIGURE 5-6 Detailed structure of the plasma membrane Although the lipid bilayer consists mainly of phospholipids, other lipids, such as cholesterol and glycolipids, are present. Peripheral proteins are loosely associated with the bilayer, whereas integral proteins are tightly bound. The integral proteins shown here are transmembrane proteins that extend through the bilayer. They have hydrophilic regions on both

sides of the bilayer, connected by a membrane-spanning a-helix. Glycolipids (carbohydrates attached to lipids) and glycoproteins (carbohydrates attached to proteins) are exposed on the extracellular surface; they play roles in cell recognition and adhesion to other cells.

close together and interacting through van der Waals interactions. In this way, unsaturated fats lower the temperature at which oil or membrane lipids solidify. Many organisms have regulatory mechanisms for maintaining cell membranes in an optimally fluid state. Some organisms compensate for temperature changes by altering the fatty acid content of their membrane lipids. When the outside temperature is cold, the membrane lipids contain relatively high proportions of unsaturated fatty acids. Some membrane lipids stabilize membrane fluidity within certain limits. One such “fluidity buffer” is cholesterol, a steroid found in animal cell membranes. A cholesterol molecule is largely hydrophobic but is slightly amphipathic because of the presence of a single hydroxyl group (see Fig. 3-15a). This hydroxyl group associates with the hydrophilic heads of the phospholipids; the hydrophobic remainder of the cholesterol molecule fits between the fatty acid hydrocarbon chains (FIG. 5-6). At low temperatures cholesterol molecules act as “spacers” between the hydrocarbon chains, restricting van der Waals interactions that would promote solidifying. Cholesterol also helps prevent the membrane from becoming weakened or unstable at higher temperatures. The reason is that the cholesterol molecules interact strongly with the portions of the hydrocarbon chains closest to the phospholipid head. This interaction restricts motion in these regions. Plant cells have steroids other than cholesterol that carry out similar functions.

Biological membranes fuse and form closed vesicles Lipid bilayers, particularly those in the liquid-crystalline state, have additional important physical properties. Bilayers tend to resist forming free ends; as a result, they are self-sealing and under most conditions spontaneously round up to form closed vesicles.

Lipid bilayers are also flexible, allowing cell membranes to change shape without breaking. Under appropriate conditions lipid bilayers fuse with other bilayers. Membrane fusion is an important cell process. When a vesicle fuses with another membrane, both membrane bilayers and their compartments become continuous. Various transport vesicles form from, and also merge with, membranes of the ER and Golgi complex, facilitating the transfer of materials from one compartment to another. A vesicle fuses with the plasma membrane when a product is secreted from the cell.

Membrane proteins include integral and peripheral proteins The two major classes of membrane proteins, integral proteins and peripheral proteins, are defined by how tightly they are associated with the lipid bilayer (FIG. 5-7). Integral membrane proteins are firmly bound to the membrane. Cell biologists usually can release them only by disrupting the bilayer with detergents. These proteins are amphipathic. Their hydrophilic regions extend out of the cell or into the cytoplasm, whereas their hydrophobic regions interact with the fatty acid tails of the membrane phospholipids. Some integral proteins do not extend all the way through the membrane. Many others, called transmembrane proteins, extend completely through the membrane. Some span the membrane only once, whereas others wind back and forth as many as 24 times. The most common kind of transmembrane protein is an α-helix (see Chapter 3) with hydrophobic amino acid side chains projecting out from the helix into the hydrophobic region of the lipid bilayer. Some proteins span the membrane in the form of rolled-up β-pleated sheets. These protein formations are barrel-shaped and form pores through which water and other substances can pass. Peripheral membrane proteins are not embedded in the lipid bilayer. They are located on the inner or outer surface of the plasma

Outside cell

Lipid bilayer Cytosol

(a) A single α-helix

(b) A protein consisting of several α-helices

(c) A rolledup β-pleated sheet

(d) Peripheral protein bound to an integral protein

FIGURE 5-7 Animated Membrane proteins Three transmembrane proteins are shown in (a), (b), and (c). The rolled-up b-pleated sheet shown in (c) forms a pore through the membrane. Water molecules and ions pass through specific types of pores (d, e). Peripheral proteins are bound to integral proteins by noncovalent interactions.

(e) Peripheral protein bound to an integral protein

RESEARCH METHOD Why Is It Used?

The freeze-fracture method is used to split the lipid bilayer apart so that its components can be analyzed.

How Is It Done? Knife edge

Ice

Cell

1

Cells are frozen in liquid nitrogen and then fractured with the edge of a knife.

2

The fracture often splits the membrane along the hydrophobic interior of the lipid bilayer.

Extracellular fluid

Bloom and Fawcett Textbook of Histology

E-face

E-face P-face

P-face Cytosol 0.1 μm

3

Two complementary fracture faces result. The inner half-membrane presents the P-face (or protoplasmic face), and the outer half-membrane presents the E-face (or external face). Integral proteins, including transmembrane proteins, are inserted through the lipid bilayer.

FIGURE 5-8 Freeze-fracture method

4

An electron microscope is used to view the interior surfaces (faces) of the two layers. In this TEM the particles (which appear as bumps) represent large integral proteins.

membrane, usually bound to exposed regions of integral proteins by noncovalent interactions. Peripheral proteins can be easily removed from the membrane without disrupting the structure of the bilayer.

Proteins are oriented asymmetrically across the bilayer One of the most remarkable demonstrations that proteins are actually embedded in the lipid bilayer comes from freeze–fracture electron microscopy, a technique that splits the membrane into halves. The researcher can literally see the two halves of the membrane from “inside out.” When cell biologists examine membranes in this way, they observe numerous particles on the fracture faces (FIG. 5-8). The particles are clearly integral membrane proteins, because researchers never see them in freeze-fractured artificial lipid bilayers. These findings profoundly influenced Singer and Nicolson in developing the fluid mosaic model. Membrane protein molecules are asymmetrically oriented. The asymmetry is produced by the highly specific way in which each

KEY POINT

protein is inserted into the bilayer. Membrane proteins that will become part of the inner surface of the plasma membrane are manufactured by free ribosomes and move to the membrane through the cytoplasm. Membrane proteins that will be associated with the cell’s outer surface are manufactured like proteins destined to be exported from the cell. As discussed in Chapter 4, proteins destined for the cell’s outer surface are initially manufactured by ribosomes on the rough ER. They pass through the ER membrane into the ER lumen, where sugars are added, making them glycoproteins. However, only a part of each protein passes through the ER membrane, so each completed protein has some regions that are located in the ER lumen and other regions that remain in the cytosol. Enzymes that attach the sugars to certain amino acids on the protein are found only in the lumen of the ER. Thus, carbohydrates can be added only to the parts of proteins located in that compartment. In FIGURE 5-9, follow from top to bottom the vesicle budding and membrane fusion events that are part of the transport process. You can see that the same region of the protein that protruded into

The orientation of a protein in the plasma membrane is determined by the pathway of its synthesis and transport.

Nucleus

Rough ER

Transport vesicle

1

Sugars are added to protein in lumen of rough ER.

2

Glycoprotein is transported to Golgi where it is further modified.

Golgi complex Plasma membrane of cell

Membrane of Golgi complex

Transport vesicle 3

Transport vesicle delivers glycoprotein to plasma membrane. Plasma membrane

Carbohydrate chain

Transport vesicle fuses with plasma membrane. Carbohydrate chain extends outward. 4

FIGURE 5-9 Synthesis and orientation of a membrane protein The surface of the rough ER membrane that faces the lumen of the rough ER also faces the lumen of the Golgi complex and vesicles. However, when a vesicle fuses with the plasma membrane, the inner surface of the vesicle becomes the extracellular surface of the plasma membrane.

the ER lumen is also transported to the lumen of the Golgi complex. There additional enzymes further modify the carbohydrate chains. Within the Golgi complex, the glycoprotein is sorted and directed to the plasma membrane. The modified region of the protein remains inside the membrane compartment of a transport vesicle as it buds from the Golgi complex. Note that when the transport vesicle fuses with the plasma membrane, the inside layer of the transport vesicle becomes the outside layer of the plasma membrane. The carbohydrate chain extends to the exterior of the cell surface. In summary, this is the sequence: sugar added to protein in ER lumen ¡ glycoprotein transported to Golgi complex, where it is further modified ¡ glycoprotein transported to plasma membrane ¡ transport vesicle fuses with plasma membrane (inside layer of transport vesicle becomes outer layer of plasma membrane)

Review ■

■

■

What molecules are responsible for the physical properties of a cell membrane? How might a transmembrane protein be positioned in a lipid bilayer? How do the hydrophilic and hydrophobic regions of the protein affect its orientation? What is the pathway used by cells to place carbohydrates on plasma membrane proteins?

5.2 OVERVIEW OF MEMBRANE PROTEIN FUNCTIONS ■ ■ LEARNING OBJECTIVE 5

KEY POINT

Cell proteins perform many functions, including transporting materials, serving as enzymes for chemical reactions, and transmitting information.

(a) Anchoring. Some membrane proteins, such as integrins, anchor the cell to the extracellular Integrin matrix; they also connect to microfilaments within the cell.

Outside cell

Inside cell

Outside cell

Inside cell

(b) Passive transport. Certain proteins form channels for selective passage of ions or molecules.

Outside cell

Inside cell

(c) Active transport. Some transport proteins pump solutes across the membrane, which requires a direct input of energy.

K+

ATP Na+

ADP

P

Outside cell

Inside cell

(d) Enzymatic activity. Many membrane-bound enzymes catalyze reactions that take place within or along the membrane surface.

Outside cell

Inside cell

(e) Signal transduction. Some receptors bind with signal molecules such as hormones and transmit information into the cell.

Outside cell

Inside bacterial cell

Summarize the functions of membrane proteins.

Why does the plasma membrane require so many different proteins? This diversity reflects the multitude of activities that take place in or on the membrane. Proteins associated with the membrane are essential for most of these activities. Generally, plasma membrane proteins fall into several broad functional categories, as shown in FIGURE 5-10. Some membrane proteins anchor the cell to its substrate. For example, integrins, proteins bound to microfilaments inside the cell, attach the cell to the extracellular matrix (Fig. 5-10a). Integrins also serve as receptors, or docking sites, for proteins of the extracellular matrix (see Fig. 4-30). Many membrane proteins are involved in the transport of molecules across the membrane. Some form channels that selectively allow the passage of specific ions or molecules (Fig. 5-10b). Other proteins form pumps that use ATP, or other energy sources, to actively transport solutes across the membrane (Fig. 5-10c). Certain membrane proteins are enzymes that catalyze reactions near the cell surface (Fig. 5-10d). In mito-

Antibody

Inside cell 1

Antigen

Inside cell 2

(f) Cell recognition. Some glycoproteins function as identification tags. For example, bacterial cells have surface proteins, or antigens, that human cells recognize as foreign.

(g) Intercellular junction. Cell adhesion proteins attach membranes of adjacent cells.

FIGURE 5-10 Some functions of membrane proteins

chondrial or chloroplast membranes, enzymes are organized in sequences that allow the organelle to efficiently regulate series of reactions in cellular respiration or photosynthesis. Some membrane proteins are receptors that receive information from other cells in the form of chemical or electrical signals. Most vertebrate cells have receptors for hormones released by endocrine glands. Information may be transmitted from proteins in the plasma membrane to the cell interior by signal transduction (discussed in Chapter 6; Fig. 5-10e). Some membrane proteins serve as identification tags that other cells recognize. For example, certain cells recognize the surface proteins, or antigens, of bacterial cells as foreign. Antigens stimulate immune defenses that destroy the bacteria (Fig. 5-10f). When certain cells recognize one another, they connect to form tissues. Some membrane proteins form junctions between adjacent cells (Fig. 5-10g). These proteins may also serve as anchoring points for networks of cytoskeletal elements. In the remaining sections of this chapter, we will discuss the functions of cell membrane proteins in transporting material into and out of the cell, and we will discuss junctions between cells. We will discuss other functions of cell membranes in many of the chapters that follow.

Review ■ ■

How do proteins function in transporting materials into the cell? What role do membrane proteins play in cell recognition?

5.3 CELL MEMBRANE STRUCTURE AND PERMEABILITY ■ ■ LEARNING OBJECTIVE 6 Describe the importance of selectively permeable membranes and compare the functions of carrier proteins and channel proteins.

A membrane is permeable to a given substance if it allows that substance to pass through and impermeable if it does not. The fluid mosaic structure of biological membranes allows them to function as selectively permeable membranes—they let some, but not all, substances pass through them. In response to varying environmental conditions or cell needs, a membrane may be a barrier to a particular substance at one time and actively promote its passage at another time. By regulating chemical traffic across its plasma membrane, a cell controls its volume and its internal ionic and molecular composition. This regulation allows the molecular composition of the cell to be quite different from that of its external environment.

Biological membranes present a barrier to polar molecules In general, biological membranes are most permeable to small nonpolar (hydrophobic) molecules. Such molecules can pass through the hydrophobic lipid bilayer. Gases such as oxygen and carbon dioxide are small, nonpolar molecules that cross the lipid bilayer rapidly. Although they are polar, water molecules are small enough to pass through gaps that occur as a fatty acid chain momentarily

moves out of the way. As a result, water molecules slowly cross the lipid bilayer. The lipid bilayer of the plasma membrane is relatively impermeable to charged ions of any size, so ions and most large polar molecules pass through the bilayer slowly. Ions are important in cell signaling and many other physiological processes. For example, many cell processes, such as muscle contraction, depend on changes in the cytoplasmic concentration of calcium ions. Glucose, amino acids, and most other compounds required in metabolism are polar molecules that also pass through the lipid bilayer slowly. This is advantageous to cells because the impermeability of the plasma membrane prevents them from diffusing out. How then do cells obtain the ions and polar molecules they require?

Transport proteins transfer molecules across membranes Systems of transport proteins that move ions, amino acids, sugars, and other needed polar molecules through membranes apparently evolved very early in the origin of cells. These transmembrane proteins have been found in all biological membranes. Two main types of membrane transport proteins are carrier proteins and channel proteins. Each type of transport protein transports a specific type of ion or molecule or a group of related substances. Carrier proteins, also called transporters, bind the ion or molecule and undergo changes in shape, resulting in movement of the molecule across the membrane. Transfer of solutes by carrier proteins located within the membrane is called carrier-mediated transport. As we will discuss, the two forms of carrier-mediated transport—facilitated diffusion and carrier-mediated active transport—differ in their capabilities and energy sources. ABC transporters make up a large, important group of carrier proteins. The acronym ABC stands for ATP-binding cassette. Found in the cell membranes of all species, ABC transporters use energy donated by ATP to transport certain ions, sugars, and polypeptides across cell membranes. Scientists have identified about 48 ABC transporters in human cells. Mutations in the genes encoding these proteins cause or contribute to many human disorders, including cystic fibrosis and certain neurological diseases. ABC transporters transport hydrophobic drugs out of the cell. This response can be a problem clinically because certain transporters remove antibiotics, antifungal drugs, and anticancer drugs. Channel proteins form tunnels, called pores, through the membrane. Many of these channels are gated, which means that they can be opened and closed. Cells regulate the passage of materials through the channels by opening and closing the gates in response to electrical changes, chemical stimuli, or mechanical stimuli. Water and specific types of ions are transported through channels. There are numerous ion channels in every membrane of every cell. Porins are transmembrane channel proteins that allow various solutes or water to pass through membranes. These channel proteins are rolled-up, barrel-shaped b-pleated sheets that form pores. Researchers Peter Agre of Johns Hopkins School of Medicine in Baltimore, Maryland, and Roderick MacKinnon of the Howard Hughes Medical Institute at Rockefeller University in New York shared the 2003 Nobel Prize in Chemistry for their work on transport proteins. Agre identified transmembrane proteins called aquaporins that function as gated water channels.

Aquaporins facilitate the rapid transport of water through the plasma membrane. About a billion water molecules per second can pass through an aquaporin! These channels are very selective and do not permit passage of ions and other small molecules. In some cells, such as those lining the kidney tubules of mammals, aquaporins respond to specific signals from hormones. Aquaporins help prevent dehydration by returning water from the kidney tubules into the blood.

Review ■ ■

■

What types of molecules pass easily through the plasma membrane? What are the two main types of transport proteins? What are their functions? What are aquaporins? What is their function?

5.4 PASSIVE TRANSPORT ■ ■ LEARNING OBJECTIVES 7 8

Contrast simple diffusion with facilitated diffusion. Define osmosis and solve simple problems involving osmosis; for example, predict whether cells will swell or shrink under various osmotic conditions.

Passive transport does not require the cell to expend metabolic energy. Many ions and small molecules move through membranes by diffusion. Two types of diffusion are simple diffusion and facilitated diffusion.

Diffusion occurs down a concentration gradient Some substances pass into or out of cells and move about within cells by diffusion, a physical process based on random motion. All atoms and molecules possess kinetic energy, or energy of motion, at temperatures above absolute zero (0 K, −273°C, or −459.4°F). Matter may exist as a solid, liquid, or gas, depending on the freedom of movement of its constituent particles (atoms, ions, or molecules). The particles of a solid are closely packed, and the forces of attraction between them let them vibrate but not move around.

1

When lump of sugar is dropped into beaker of pure water, sugar molecules begin to dissolve and diffuse through water.

FIGURE 5-11 Diffusion

2

Sugar molecules continue to dissolve and diffuse through water.

In a liquid the particles are farther apart; the intermolecular attractions are weaker, and the particles move about with considerable freedom. In a gas the particles are so far apart that intermolecular forces are negligible; molecular movement is restricted only by the walls of the container that encloses the gas. Atoms and molecules in liquids and gases move in a kind of “random walk,” changing directions as they collide. Although the movement of individual particles is undirected and unpredictable, we can nevertheless make predictions about the behavior of groups of particles. If the particles are not evenly distributed, then at least two regions exist: one with a higher concentration of particles and the other with a lower concentration. Such a difference in the concentration of a substance from one place to another establishes a concentration gradient. In diffusion, the random motion of particles results in their net movement “down” their own concentration gradient, from the region of higher concentration to the one of lower concentration. This does not mean individual particles are prohibited from moving “against” the gradient. However, because there are initially more particles in the region of high concentration, it logically follows that more particles move randomly from there into the lowconcentration region than the reverse (FIG. 5-11). Thus, if a membrane is permeable to a substance, there is net movement from the side of the membrane where it is more highly concentrated to the side where it is less concentrated. Such a gradient across the membrane is a form of stored energy. Stored energy is potential energy, which is the capacity to do work as a result of position or state. The stored energy of the concentration gradient is released when ions or molecules move from a region of high concentration to one of low concentration. For this reason, movement down a concentration gradient is spontaneous. (Forms of energy and spontaneous processes are discussed in greater detail in Chapter 7.) Diffusion occurs rapidly over very short distances. The rate of diffusion is determined by the movement of the particles, which in turn is a function of their size and shape, their electric charges, and the temperature. As the temperature rises, particles move faster and the rate of diffusion increases. Particles of different substances in a mixture diffuse independently of one another. Diffusion moves solutes toward a state of equilibrium. If particles are not added to or removed from the system, a state of dynamic equilibrium is reached. In this condition, the particles are uniformly distributed and there is no net change in the system. Particles continue to move back and forth across the membrane, but they move at equal rates and in both directions. In organisms, equilibrium is rarely attained. For example, hu3 Eventually, sugar man cells continually produce carmolecules become distributed randomly bon dioxide as sugars and other throughout water. molecules are metabolized during aerobic respiration. Carbon dioxide readily diffuses across the plasma

membrane but then is rapidly removed by the blood. This limits the opportunity for the molecules to re-enter the cell, so a sharp concentration gradient of carbon dioxide molecules always exists across the plasma membrane. In simple diffusion through a biological membrane, small, nonpolar (uncharged) solute molecules move directly through the membrane down their concentration gradient. Oxygen and carbon dioxide can rapidly diffuse through the membrane. The rate of simple diffusion is directly related to the concentration of the solute; the more concentrated the solute, the more rapid the diffusion.

Osmosis is diffusion of water across a selectively permeable membrane

Pressure applied to piston to resist upward movement

Water plus solute

Pure water

Selectively permeable membrane

Osmosis is a special kind of diffusion that involves the net movement of water (the principal solvent in biological systems) through a selectively permeable membrane from a region of higher conMolecule of solute Water centration to a region of lower concentration. Water molecules molecule pass freely in both directions, but as in all types of diffusion, net movement is from the region where the water molecules are more concentrated to the region where they are less concentrated. Most solute molecules (such as sugar and salt) cannot diffuse freely through the selectively permeable membranes of the cell. FIGURE 5-12 Animated Osmosis The principles involved in osmosis can be illustrated using an The U-tube contains pure water on the right and water plus a solute on the left, separated by a selectively permeable membrane. Water molecules apparatus called a U-tube (FIG. 5-12). The U-tube is divided into cross the membrane in both directions (blue arrows). Solute molecules cantwo sections by a selectively permeable membrane that allows not cross (red arrows). The fluid level would normally rise on the left and fall solvent (water) molecules to pass freely but excludes solute moleon the right because net movement of water would be to the left. However, cules. A water/solute solution is placed on one side, and pure water the piston prevents the water from rising. The force that must be exerted by is placed on the other. The side containing the solute has a lower the piston to prevent the rise in fluid level is equal to the osmotic pressure of the solution. effective concentration of water than the pure water side does. The reason is that the solute particles, which are charged (ionic) or polar, interact with the partial electric charges on the polar water by the piston to prevent the rise of fluid on that side of the tube. A molecules. Many of the water molecules are thus “bound up” and solution with a high solute concentration has a low effective water no longer free to diffuse across the membrane. concentration and a high osmotic pressure; conversely, a solution Because of the difference in effective water concentration, with a low solute concentration has a high effective water concenthere is net movement of water molecules from the pure water side tration and a low osmotic pressure. (with a high effective concentration of water) to the water/ solute side (with a lower effective concentration of water). As a result, Two solutions may be isotonic the fluid level drops on the pure water side and rises on the water/ Salts, sugars, and other substances are dissolved in the fluid comsolute side. Because the solute molecules do not diffuse across the partment of every cell. These solutes give the cytosol a specific membrane, equilibrium is never attained. Net movement of waosmotic pressure. TABLE 5-1 summarizes the movement of water ter continues, and the fluid level rises on the side containing the into and out of a solution (or cell) depending on relative solute solute. The weight of the rising column of fluid eventually exerts concentrations. enough pressure to stop further changes in fluid levels, although When a cell is placed in a fluid with exactly the same osmotic water molecules continue to pass through the selectively permepressure, no net movement of water molecules occurs, either into able membrane in both directions. We define the osmotic pressure of a solution as the pressure that must be exerted on the side of a selectively permeable TABLE 5-1 Osmotic Terminology membrane containing the higher concenSolute Solute Direction of tration of solute to prevent the diffusion of Concentration Concentration Net Movement water (by osmosis) from the side containin Solution A in Solution B Tonicity of Water ing the lower solute concentration. In the U-tube example, you could measure the Greater Less A hypertonic to B; B hypotonic to A B to A osmotic pressure by inserting a piston on Less Greater B hypertonic to A; A hypotonic to B A to B the water/solute side of the tube and meaEqual Equal A and B are isotonic to each other No net movement suring how much pressure must be exerted

Outside cell

Inside cell

Outside cell

Inside cell

Outside cell

Inside cell

H2O molecules Solute molecules

Net water movement out of the cell

Net water movement into the cell

10 μm

Courtesy of Dr. R. F. Baker, Emeritus/ USC School of Medicine

No net water movement

(a) Isotonic solution. When a cell is placed in an isotonic solution, water molecules pass in and out of the cell, but the net movement of water molecules is zero.

(b) Hypertonic solution. When a cell is placed in a hypertonic solution, there is a net movement of water molecules out of the cell (blue arrow ). The cell becomes dehydrated and shrunken.

(c) Hypotonic solution. When a cell is placed in a hypotonic solution, the net movement of water molecules into the cell (blue arrow) causes the cell to swell or even burst.

FIGURE 5-13 Animated The responses of animal cells to osmotic pressure differences

or out of the cell. The cell neither swells nor shrinks. Such a fluid is of equal solute concentration, or isotonic, to the fluid within the cell. Normally, your blood plasma (the fluid component of blood) and all your other body fluids are isotonic to your cells; they contain a concentration of water equal to that in the cells. A solution of 0.9% sodium chloride (sometimes called physiological saline) is isotonic to the cells of humans and other mammals. Human red blood cells placed in 0.9% sodium chloride neither shrink nor swell (FIG. 5-13a). One solution may be hypertonic and the other hypotonic If the surrounding fluid has a concentration of dissolved substances greater than the concentration within the cell, the fluid has a higher osmotic pressure than the cell and is said to be hypertonic to the cell. Because a hypertonic solution has a lower effective water concentration, a cell placed in such a solution shrinks as it loses water

by osmosis. Human red blood cells placed in a solution of 1.3% sodium chloride shrivel (FIG. 5-13b). If the surrounding fluid contains a lower concentration of dissolved materials than does the cell, the fluid has a lower osmotic pressure and is said to be hypotonic to the cell; water then enters the cell and causes it to swell. Red blood cells placed in a solution of 0.6% sodium chloride gain water, swell (FIG. 5-13c), and may eventually burst. Many cells that normally live in hypotonic environments have adaptations to prevent excessive water accumulation. For example, Paramecium and certain other ciliates (members of the supergroup Chromalveolates) have contractile vacuoles that expel excess water (see Fig. 26-8). Turgor pressure is the internal hydrostatic pressure usually present in walled cells The cells of most prokaryotes, algae, plants, and fungi have relatively rigid cell walls. These cells can withstand, without bursting,

Cengage

Plasma membrane

Nucleus Vacuole

Vacuole Vacuolar membrane (tonoplast) Cytoplasm

(a) In hypotonic surroundings, the vacuole of a plant cell fills with water, but the rigid cell walls prevent the cell from expanding. The cells of this healthy begonia plant are turgid.

Plasma membrane

(b) When the begonia plant is exposed to a hypertonic solution, its cells become plasmolyzed as they lose water.

(c) The plant wilts and eventually dies.

FIGURE 5-14 Animated Turgor pressure and plasmolysis

an external medium that is very dilute, containing only a very low concentration of solutes. Because of the substances dissolved in the cytoplasm, the cells are hypertonic to the outside medium (conversely, the outside medium is hypotonic to the cytoplasm). Water moves into the cells by osmosis, filling their central vacuoles and distending the cells. The cells swell, building up turgor pressure against the rigid cell walls (FIG. 5-14a). The cell walls stretch only slightly, and a steady state is reached when their resistance to stretching prevents any further increase in cell size and thereby halts the net movement of water molecules into the cells. (Of course, molecules continue to move back and forth across the plasma membrane.) Turgor pressure in the cells is an important factor in supporting the body of nonwoody plants. If a cell that has a cell wall is placed in a hypertonic medium, the cell loses water to its surroundings. Its contents shrink, and the plasma membrane separates from the cell wall, a process known as plasmolysis (FIGS. 5-14b and 14c). Plasmolysis occurs in plants

when the soil or water around them contains high concentrations of salts or fertilizers. It also explains why lettuce becomes limp in a salty salad dressing and why a picked flower wilts from lack of water.

Facilitated diffusion occurs down a concentration gradient In all processes in which substances move across membranes by diffusion, the net transfer of those molecules from one side to the other occurs as a result of a concentration gradient. We have seen that small, uncharged (nonpolar) solute molecules, such as oxygen and carbon dioxide, move directly through the membrane down their concentration gradient by simple diffusion. In facilitated diffusion, a specific transport protein makes the membrane permeable to a particular solute, such as a specific ion or polar molecule. A specific solute can be transported from inside the cell

Outside cell

K+

K+

K+ K+

gradients (FIG. 5-15). (As we will discuss, because ions are charged particles, these gradients are electrochemical gradients.) These ion channels are referred to as gated channels because they can open and close. As many as 100 million ions per second can pass through an open ion channel! Channels can facilitate transport only down a concentration gradient. They cannot actively transport ions from a region of lower concentration to a region of higher concentration.

K+

K+

K+

K+ Cytosol

FIGURE 5-15 Facilitated diffusion of potassium ions In response to an electrical stimulus, the gate of the potassium ion channel opens, allowing potassium to diffuse out of the cell.

to the outside or from the outside to the inside, but net movement is always from a region of higher solute concentration to a region of lower concentration. Channel proteins and carrier proteins facilitate diffusion by different mechanisms. Channel proteins form hydrophilic channels through membranes Some channel proteins are porins that form rather large tunnels through which water and solutes pass. However, most channel proteins form narrow channels that transport specific ions down their

KEY POINT

Carrier proteins undergo a change in shape Transport of solutes through carrier proteins is slower than through channel proteins. The carrier protein binds with one or more solute molecules on one side of the membrane. The protein then undergoes a conformational change (change in shape) that moves the solute to the other side of the membrane. As an example of facilitated diffusion by a carrier protein, let us consider glucose transport. A carrier protein known as glucose transporter 1, or GLUT 1, transports glucose into red blood cells (FIG. 5-16). The concentration of glucose is higher in the blood plasma than in red blood cells, so glucose diffuses down its concentration gradient into these blood cells. The GLUT 1 transporter facilitates glucose diffusion, allowing glucose to enter the cell about 50,000 times as rapidly as it could by simple diffusion. Red blood cells keep the internal concentration of glucose low by immediately adding a phosphate group to entering glucose molecules, converting them to highly charged glucose phosphates that cannot pass back through the membrane. Because glucose phosphate is a different molecule, it does not contribute to the glucose concentration gradient. Thus, a steep concentration gradient for glucose is continually maintained, and glucose rapidly diffuses into the

Facilitated diffusion requires the potential energy of a concentration gradient.

Outside cell Glucose High concentration of glucose

Low concentration of glucose

Glucose transporter (GLUT 1)

Cytosol 1

Glucose binds to GLUT 1.

2

GLUT 1 changes shape and glucose is released inside cell.

FIGURE 5-16 Animated Facilitated diffusion of glucose molecules

3

GLUT 1 returns to its original shape.

cell, only to be immediately changed to the phosphorylated form. Facilitated diffusion is powered by the concentration gradient. Researchers have studied facilitated diffusion of glucose using liposomes, artificial vesicles surrounded by phospholipid bilayers. The phospholipid membrane of a liposome does not allow the passage of glucose unless a glucose transporter is present in the liposome membrane. Glucose transporters and similar carrier proteins temporarily bind to the molecules they transport. This mechanism appears to be similar to the way an enzyme binds with its substrate, the molecule on which it acts (discussed in Chapter 7). In addition, as in enzyme action, binding apparently changes the shape of the carrier protein. This change allows the glucose molecule to be released on the inside of the cell. According to this model, when the glucose is released into the cytoplasm, the carrier protein reverts to its original shape and is available to bind another glucose molecule on the outside of the cell. Another similarity to enzyme action is that carrier proteins become saturated when there is a high concentration of the molecule being transported. This saturation may occur because a finite number of carrier proteins are available and they operate at a defined maximum rate. When the concentration of solute molecules to be transported reaches a certain level, all the carrier proteins are working at their maximum rate. It is a common misconception that diffusion, whether simple or facilitated, is somehow “free of cost” and that only active transport mechanisms require energy. Because diffusion always involves the net movement of a substance down its concentration gradient, we say that the concentration gradient “powers” the process. However, energy is required to do the work of establishing and maintaining the gradient. In our example of facilitated diffusion of glucose, the cell maintains a steep concentration gradient (high outside, low inside) by phosphorylating the glucose molecules once they enter the cell. One ATP molecule is spent for every glucose molecule phosphorylated, and there are additional costs, such as the energy required to make the enzymes that carry out the reaction.

Review ■

■

■

What would happen if a plant cell were placed in an isotonic solution? a hypertonic environment? a hypotonic environment? How would you modify your predictions for an animal cell? What is the immediate source of energy for simple diffusion? For facilitated diffusion? In what direction do particles move along their concentration gradient? Would your answers be different for facilitated diffusion compared with simple diffusion?

5.5 ACTIVE TRANSPORT ■ ■ LEARNING OBJECTIVE 9 Describe active transport, including cotransport.

Although adequate amounts of a few substances move across cell membranes by diffusion, cells must actively transport many solutes against a concentration gradient. The reason is that cells

require many substances in higher concentrations than their concentration outside the cell. Both diffusion and active transport require energy. The energy for diffusion is provided by a concentration gradient for the substance being transported. Active transport requires the cell to expend metabolic energy directly to power the process. An active transport system can pump materials from a region of low concentration to a region of high concentration. The energy stored in the concentration gradient not only is unavailable to the system but actually works against it. For this reason, the cell needs some other source of energy. In many cases, cells use ATP energy directly. However, active transport may be coupled to ATP indirectly. In indirect active transport, a concentration gradient provides energy for the cotransport of some other substance, such as an ion.

Active transport systems “pump” substances against their concentration gradients One of the most striking examples of an active transport mechanism is the sodium–potassium pump found in all animal cells (FIG. 5-17). The pump is an ABC transporter, a specific carrier protein in the plasma membrane. This transporter uses energy from ATP to pump sodium ions out of the cell and potassium ions into the cell. The exchange is unequal: usually only two potassium ions are imported for every three sodium ions exported. Because these particular concentration gradients involve ions, an electrical potential (separation of electric charges) is generated across the membrane; that is, the membrane is polarized. Both sodium and potassium ions are positively charged, but because there are fewer potassium ions inside relative to the sodium ions outside, the inside of the cell is negatively charged relative to the outside. The unequal distribution of ions establishes an electrical gradient that drives ions across the plasma membrane. Sodium–potassium pumps help maintain a separation of charges across the plasma membrane. This separation is called a membrane potential. Because there is both an electric charge difference and a concentration difference on the two sides of the membrane, the gradient is called an electrochemical gradient. Such gradients store energy that is used to drive other transport systems. So important is the electrochemical gradient produced by these pumps that some cells (such as nerve cells) expend more than 25% of their total available energy just to power this one transport system. Sodium–potassium pumps (as well as all other ATP-driven pumps) are transmembrane proteins that extend entirely through the membrane. By undergoing a series of conformational changes, the pumps exchange sodium for potassium across the plasma membrane. Unlike what occurs in facilitated diffusion, at least one of the conformational changes in the pump cycle requires energy, which is provided by ATP. The shape of the pump protein changes as a phosphate group from ATP first binds to it and is subsequently removed later in the pump cycle. The use of electrochemical potentials for energy storage is not confined to the plasma membranes of animal cells. Cells of bacteria, fungi, and plants use carrier proteins, known as proton pumps, to actively transport hydrogen ions (which are protons) out of the cell. These ATP-driven membrane pumps transfer protons

KEY POINT

The sodium–potassium pump is a carrier protein that maintains an electrochemical gradient across the plasma membrane.

Higher

Na+ Na+

Lower

Outside cell

Na+

Potassium concentration gradient

Sodium concentration gradient

Active transport channel

ADP + P

ATP

K+ +

K

Cytosol

Lower

Higher

(a) The sodium–potassium pump is a carrier protein that requires energy from ATP. In each complete pumping cycle, the energy of one molecule of ATP is used to export three sodium ions (Na+) and import two potassium ions (K+).

Na+

Na+

Na+

Na+

ATP

Na+

P Na+ +

K

P

ADP 2

+

K

Phosphate group is transferred from ATP to carrier protein.

3

Phosphorylation causes carrier protein to change shape, releasing 3 Na+ outside cell. P

Na+

1

Na+

Na+

Three Na+ bind to carrier protein.

4 K+ K+

+

K

K+

P

5 Phosphate is released. Phosphate release causes carrier protein to return to its original shape. Two K+ ions are released inside cell. (b) Follow the steps illustrating a model of active transport by the sodium–potassium pump. 6

FIGURE 5-17 Animated A model for the pumping cycle of the sodium–potassium pump

Two K+ bind to carrier protein.

from the cytosol to the outside (FIG. 5-18). Removal of positively charged protons from the cytoplasm of these cells results in a large difference in the concentration of protons between the outside and inside of the cell. The outside of the cells is positively charged relative to the inside of the plasma membrane. The energy stored in these electrochemical gradients can be used to do many kinds of cell work. Other proton pumps are used in “reverse” to synthesize ATP. Bacteria, mitochondria, and chloroplasts use energy from food or sunlight to establish proton concentration gradients (discussed in Chapters 8 and 9). When the protons diffuse through the proton carriers from a region of high proton concentration to one of low concentration, ATP is synthesized. These electrochemical gradients form the basis for the major energy conversion systems in virtually all cells. Ion pumps have other important roles. For example, they are instrumental in the ability of an animal cell to equalize the osmotic pressures of its cytoplasm and its external environment. If an animal cell does not control its internal osmotic pressure, its contents become hypertonic relative to the exterior. Water enters by osmosis, causing the cell to swell and possibly burst (see Fig. 5-13c). By controlling the ion distribution across the membrane, the cell indirectly controls the movement of water, because when ions are pumped out of the cell, water leaves by osmosis.

Carrier proteins can transport one or two solutes You may have noticed that some carrier proteins, such as proton pumps, transport one type of substance in one direction. These carrier proteins are called uniporters. Other carrier proteins, symporters, move two types of substances in one direction. For example, a specific carrier protein transports both sodium and glucose into the cell. Still other carrier proteins, antiporters, move two substances in opposite directions. Sodium–potassium pumps transport sodium ions out of the cell and potassium ions into the cell. Both symporters and antiporters cotransport solutes.

Cotransport systems indirectly provide energy for active transport A cotransport system moves solutes across a membrane by indirect active transport. Two solutes are transported at the same time. The movement of one solute down its concentration gradient provides energy for transport of some other solute up its concentration gradient. However, an energy source such as ATP is required to power the pump that produces the concentration gradient. Sodium–potassium pumps (and other pumps) generate electrochemical concentration gradients. Sodium is pumped out of the cell and then diffuses back in by moving down its concentration gradient. This process generates sufficient energy to power the active transport of other essential substances. In these systems, a carrier protein cotransports a solute against its concentration gradient, while sodium, potassium, or hydrogen ions move down their gradient. Energy from ATP produces the ion gradient. Then the energy of this gradient drives the active transport of a required substance, such as glucose, against its gradient.

H+

H+ Outside cell

+

– Cytosol

H+

H+

+

+

–

– ATP ADP

H+

+

+

– –

H+

FIGURE 5-18 A model of a proton pump Proton pumps use the energy of ATP to transport protons (hydrogen ions) across membranes. The energy of the electrochemical gradient established can then be used for other processes.

We have seen how glucose can be moved into the cell by facilitated diffusion. Glucose can also be cotransported into the cell. The sodium concentration inside the cell is kept low by the ATPrequiring sodium–potassium pumps that actively transport sodium ions out of the cell. In glucose cotransport, a carrier protein transports both sodium and glucose (FIG. 5-19). As sodium moves into the cell along its concentration gradient, the carrier protein captures the energy released and uses it to transport glucose into the cell. Thus, this indirect active transport system for glucose is “driven” by the cotransport of sodium.

Review ■ ■

What is the energy source for active transport? What is the energy source for cotransport?

5.6 EXOCYTOSIS AND ENDOCYTOSIS ■ ■ LEARNING OBJECTIVE 10 Compare exocytotic and endocytotic transport mechanisms.

Individual molecules and ions pass through the plasma membrane by simple and facilitated diffusion and by carrier-mediated active transport. Some larger materials, such as large molecules, particles of food, and even small cells, are also moved into or out of cells. They are transported by exocytosis and endocytosis. Like active transport, these processes require cells to expend energy directly.

In exocytosis, vesicles export large molecules In exocytosis, a cell ejects waste products, or specific products of secretion such as hormones, by the fusion of a vesicle with the

KEY POINT

A carrier protein transports sodium ions down their concentration gradient and uses that energy to cotransport glucose molecules against their concentration gradient.

Outside cell +

+

Carrier protein + +

+

+

+ +

+ +

+

+

+

+

+

+

–

Glucose

Na+

+

+

+

+ +

+

+

+

+ +

+ +

+

+

+

+

–

–

–

–

Lipid bilayer

–

–

Glucose concentration gradient

–

+ + +

+

+

Cytosol 1

Sodium ions and glucose bind to carrier protein.

2

Carrier protein changes shape and releases sodium ions and glucose inside cell.

FIGURE 5-19 A model for the cotransport of glucose and sodium ions Note that this carrier protein is a symporter.

2

3

1

Vesicle approaches plasma membrane,

2

fuses with it, and

Bloom and Fawcett Textbook of Histology

1

0.25 μm

FIGURE 5-20 Exocytosis 3

releases its contents outside cell.

The TEM shows exocytosis of the protein components of milk by a mammary gland cell.

Vacuole Lysosomes

Folds of plasma membrane surround particle to be ingested, forming small vacuole around it.

Bloom and Fawcett Textbook of Histology

Nucleus

2

Vacuole then pinches off inside cell.

Ingested bacteria

3

Lysosomes fuse with vacuole and pour potent hydrolytic enzymes onto ingested material.

2.5 μm

1

Lysosome

Lysosome

Glycogen (stored nutrients)

FIGURE 5-21 Animated Phagocytosis Bacteria Lysosomes

Large vacuole

plasma membrane (FIG. 5-20). As the contents of the vesicle are released from the cell, the membrane of the secretory vesicle is incorporated into the plasma membrane. This is the primary mechanism by which plasma membranes grow larger.

In endocytosis, the cell imports materials

In this type of endocytosis, a cell ingests relatively large solid particles. The white blood cell (a neutrophil) shown in the TEM is phagocytizing bacteria. The vacuoles contain bacteria that have already been ingested. Lysosomes contain digestive enzymes that break down the ingested material. Other bacteria are visible outside the cell.

contents of these vesicles are slowly transferred into the cytosol, the vesicles become progressively smaller. In a third type of endocytosis, receptor-mediated endocytosis, specific molecules combine with receptor proteins in the plasma membrane. Receptor-mediated endocytosis is the main mechanism by which eukaryotic cells take in macromolecules. As an example, let us look at how mammalian cells take up cholesterol from the blood. Cells use cholesterol as a component of cell membranes and as a precursor of steroid hormones. Cholesterol is transported in the blood as part of particles called low-density lipoproteins (LDLs; popularly known as “bad cholesterol”).

In endocytosis, materials are taken into the cell. Several types of endocytotic mechanisms operate in biological systems, including phagocytosis, pinocytosis, and receptor-mediated endocytosis. In phagocytosis (literally, “cell eating”), the cell ingests large solid particles Microvilli such as food or bacteria (FIG. 5-21). Certain protists ingest food by phagocytosis. Some types of vertebrate cells, including certain white blood cells, ingest bacteria and other particles by phagocytosis. During ingestion, folds of the plasma membrane enclose the cell or particle. When the membrane has encircled the particle, the membrane fuses at the point of contact, forming a vacuole. The vacuole may then fuse with lysosomes, Cytosol which degrade the ingested material. In pinocytosis (“cell drinking”), the 1 Tiny droplets of fluid 2 These pinch off into cell takes in dissolved materials (FIG. 5-22). are trapped by folds cytosol as small of plasma membrane. fluid-filled vesicles. Tiny droplets of fluid are trapped by folds in the plasma membrane, which pinch off into FIGURE 5-22 Pinocytosis, or “cell drinking” the cytosol as tiny vesicles. As the liquid

Pinocytotic vesicle

3

Contents of these vesicles are then slowly transferred to cytosol.

Much of the receptor-mediated endocytosis pathway was detailed through studies at the University of Texas Health Science Center by Michael Brown and Joseph Goldstein on the LDL receptor. These researchers were awarded the Nobel Prize in Physiology or Medicine in 1985 for their pioneering work. Their findings have important medical implications because cholesterol that remains in the blood instead of entering the cells can be deposited in the artery walls, which increases the risk of cardiovascular disease. When it needs cholesterol, the cell makes LDL receptors. The receptors are concentrated in coated pits, depressed regions

KEY POINT

In receptor-mediated endocytosis, specific macromolecules bind to receptor proteins, accumulate in coated pits, and enter the cell in clathrin-coated vesicles. Plasma membrane

1

2

Cytosol 2

LDL attaches to specific receptors in coated pits on plasma membrane. Endocytosis results in formation of a coated vesicle in cytosol. Seconds later coat is removed.

3

Uncoated vesicle fuses with endosome.

4

Receptors are returned to plasma membrane and recycled.

5

on the cytoplasmic surface of the plasma membrane. Each pit is coated by a layer of a protein, called clathrin, found just below the plasma membrane. A molecule that binds specifically to a receptor is called a ligand. In this case, LDL is the ligand. After the LDL binds with a receptor, the coated pit forms a coated vesicle by endocytosis. FIGURE 5-23 shows the uptake of an LDL particle. Seconds after the vesicle moves into the cytoplasm, the coating dissociates from it, leaving an uncoated vesicle. The vesicles fuse with small compartments called endosomes. The LDL and LDL receptors sep-

Uncoated vesicle

Coated pit LDL particle

1

LDL receptor

Clathrin recycled

Endosome

Primary

3

lysosome

Clathrin

Vesicle containing LDL particles fuses with a lysosome, forming a secondary lysosome. Hydrolytic enzymes then digest cholesterol from LDL particles for use by cell.

Endosome

5

Secondary lysosome 4

Free cholesterol

(a) Uptake of low-density lipoprotein (LDL) particles, which transport cholesterol in the blood.

From M.M. Perry and A.B. Gilbert, Journal of Cell Science 39: 257–272, 1979. © 1979 The Company of Biologists Ltd.

(b) This series of TEMs shows the formation of a coated vesicle from a coated pit.

FIGURE 5-23 Animated Receptor-mediated endocytosis

0.25 μm

ligand molecules bind to receptors in coated pits of plasma membrane ¡ coated vesicle forms by endocytosis ¡ coating detaches from vesicle ¡ uncoated vesicle fuses with endosome ¡ ligands separate from receptors which are recycled; endosome fuses with primary lysosome, forming secondary lysosome ¡ contents of secondary lysosome are digested and released into the cytosol

The recycling of LDL receptors to the plasma membrane through vesicles causes a problem common to all cells that use endocytotic and exocytotic mechanisms: the plasma membrane changes size as the vesicles bud off from it or fuse with it. A type of phagocytic cell known as a macrophage, for example, ingests the equivalent of its entire plasma membrane in about 30 minutes, requiring an equivalent amount of recycling or new membrane synthesis for the cell to maintain its surface area. On the other hand, cells that are constantly involved in secretion must return an equivalent amount of membrane to the interior of the cell for each vesicle that fuses with the plasma membrane; if not, the cell surface would continue to expand even though the growth of the cell itself may be arrested.

choring junctions. These junctions do not prevent the passage of materials between adjacent cells. Two common types of anchoring junctions are desmosomes and adhering junctions. Desmosomes are points of attachment between cells (FIG. 5-24). They hold cells together at one point as a rivet or a spot weld does. Desmosomes allow cells to form strong sheets, and substances still pass freely through the spaces between the plasma membranes. Each desmosome is made up of regions of dense material associated with the cytosolic sides of the two plasma membranes, plus protein filaments that cross the narrow intercellular space between them. Desmosomes are anchored to systems of

Bloom and Fawcett Textbook of Histology

arate, and the receptors are transported to the plasma membrane, where they are recycled. LDL is transferred to a lysosome, where it is broken down. Cholesterol is released into the cytosol for use by the cell. A simplified summary of receptor-mediated endocytosis follows:

Plasma membranes

0.25 μm

Review ■ ■ ■

How are exocytosis and endocytosis similar? How are the processes of phagocytosis and pinocytosis different? What is the sequence of events in receptor-mediated endocytosis?

5.7 CELL JUNCTIONS ■ ■ LEARNING OBJECTIVE 11 Compare the structures and functions of anchoring junctions, tight junctions, gap junctions, and plasmodesmata.

Intercellular space Intermediate filaments Desmosome

Protein filaments Disc of dense protein material

Cells in close contact with one another typically develop specialized intercellular junctions. These structures may allow neighboring cells to form strong connections with one another, prevent the passage of materials, or establish rapid communication between adjacent cells. Several types of junctions connect animal cells, including anchoring junctions, tight junctions, and gap junctions. Plant cells are connected by plasmodesmata.

Anchoring junctions connect cells of an epithelial sheet Adjacent epithelial cells, such as those found in the outer layer of the mammalian skin, are so tightly bound to each other by anchoring junctions that strong mechanical forces are required to separate them. Cadherins, transmembrane proteins shown in the chapter opening photograph, are important components of an-

Cell 1

Cell 2

FIGURE 5-24 Desmosomes The dense structure in the TEM is a desmosome. Each desmosome consists of a pair of buttonlike discs associated with the plasma membranes of adjacent cells, plus the intercellular protein filaments that connect them. Intermediate filaments in the cells are attached to the discs and are connected to other desmosomes.

intermediate filaments inside the cells. Thus, the intermediate filament networks of adjacent cells are connected. As a result, mechanical stresses are distributed throughout the tissue. Adhering junctions cement cells together. Cadherins form a continuous adhesion belt around each cell, binding the cell to neighboring cells. These junctions connect to microfilaments of the cytoskeleton. The cadherins of adhering junctions are a potential path for signals from the outside environment to be transmitted to the cytoplasm.

Tight junctions seal off intercellular spaces between some animal cells Tight junctions are literally areas of tight connections between the membranes of adjacent cells. These connections are so tight

KEY POINT

that no space remains between the cells and substances cannot leak between them. TEMs of tight junctions show that in the region of the junction the plasma membranes of the two cells are held together by proteins in actual contact with each other. However, as shown in FIGURE 5-25, tight junctions are located intermittently. The plasma membranes of the two cells are not fused over their entire surface. Cells connected by tight junctions seal off body cavities. For example, tight junctions between cells lining the intestine prevent substances in the intestine from passing between the cells and directly entering the blood. The sheet of cells thus acts as a selective barrier. Food substances must be transported across the plasma membranes and through the intestinal cells before they enter the blood. This arrangement helps prevent toxins and other unwanted materials from entering the blood and also prevents nutrients

Tight junctions prevent the passage of materials through spaces between cells.

Microvillus

Lumen of the intestine

Tight junction Intercellular space

Cell 1

Cell 2

G. E. Palade

Rows of tight junction proteins

Plasma membranes

Intercellular space

0.1 Nm

(a) This TEM shows points of fusion between plasma membranes of adjacent cells lining the intestine. One tight junction is marked by the box.

FIGURE 5-25 Animated Tight junctions

(b) A tight junction is formed by linkages between rows of proteins of adjacent cells. These proteins seal off the intercellular space, preventing passage of materials through spaces between cells.

from leaking out of the intestine. Tight junctions are also present between the cells that line capillaries in the brain. They form the blood–brain barrier, which prevents many substances in the blood from passing into the brain.

Gap junctions allow the transfer of small molecules and ions A gap junction is like a desmosome in that it bridges the space between cells; however, the space it spans is somewhat narrower (FIG. 5-26). Gap junctions also differ in that they are communicating junctions. They not only connect the plasma membranes but also contain channels connecting the cytoplasm of adjacent cells. Gap junctions are composed of connexin, an integral membrane protein. Groups of six connexin molecules cluster to form a cylinder that spans the plasma membrane. The connexin cylinders on adjacent cells become tightly joined. The two cylinders form a channel, about 1.5 nm in diameter. Small inorganic molecules (such as ions) and some regulatory molecules (such as cyclic AMP, which is illustrated in Figure 3-25) pass through the channels, but larger molecules are excluded. When a marker substance is in-

Plasmodesmata allow certain molecules and ions to move between plant cells Because plant cells have walls, they do not need desmosomes for strength. Plant cells have connections that are functionally equivalent to the gap junctions of some animal cells. Plasmodesmata

Gap junctions allow the transfer of small molecules and ions between adjacent cells.

E. Anderson et al. Journal of Morphology 156: 339–366, 1978. Reprinted with permission of Wiley–Liss, Inc., a subsidiary of John Wiley & Sons, Inc.

KEY POINT

jected into one of a group of cells connected by gap junctions, the marker passes rapidly into the adjacent cells but does not enter the space between the cells. Gap junctions provide for rapid chemical and electrical communication between cells. Cells control the passage of materials through gap junctions by opening and closing the channels (see Fig. 5-26d). Cells in the pancreas, for example, are linked by gap junctions in such a way that if one of a group of cells is stimulated to secrete insulin, the signal is passed through the junctions to the other cells in the cluster. This mechanism ensures a coordinated response to the initial signal. Gap junctions allow some nerve cells to be electrically coupled. Cardiac muscle cells are linked by gap junctions that permit the flow of ions necessary to synchronize contractions of the heart.

0.25 Nm

Bloom and Fawcett Textbook of Histology

(c) A freeze–fracture replica of the P-face of a gap junction between two ovarian cells of a mouse. Each particle corresponds to a connexin cylinder.

Closed

0.1 Nm

(a) A TEM of a gap junction.

(b) The two plasma membranes contain cylinders composed of six connexin molecules. Two cylinders from opposite membranes join to form a channel connecting the cytoplasmic compartments of the two cells.

(d) This model shows how a gap junction pore might open and close.

FIGURE 5-26 Animated Gap junctions The model of a gap junction shown in (b) is based on electron microscopic and X-ray diffraction data.

Open

(sing., plasmodesma) are channels 20 to 40 nm wide that pass through the cell walls of adjacent plant cells, connecting the cytoplasm of neighboring cells (FIG. 5-27). The plasma membranes of adjacent cells are continuous with one another through the plasmodesmata. Most plasmodesmata contain a narrow cylindrical structure, called the desmotubule, which runs through the channel and connects the smooth ER of the two adjacent cells. Plasmodesmata generally allow molecules and ions, but not organelles, to pass through the openings from cell to cell. The movement of ions through the plasmodesmata allows for a very slow type of electrical signaling in plants. Whereas the channels of gap junctions have a fixed diameter, plants cells can dilate the plasmodesmata channels. Certain proteins and RNA can pass through plasmodesmata. Some plant viruses spread infection by passing through these junctions.

KEY POINT

Cell walls

Desmotubule Plasma membrane

Smooth ER

How are desmosomes and tight junctions functionally similar? How do they differ? What is the justification for considering gap junctions and plasmodesmata to be functionally similar? How do they differ structurally?

■

Cell 1

Plasmodesmata

Review ■

Most plant cells have plasmodesmata that connect the cytoplasm of adjacent cells.

Cell 2

FIGURE 5-27 Plasmodesmata Cytoplasmic channels through the cell walls of adjacent plant cells allow passage of water, ions, and small molecules. The channels are lined with the fused plasma membranes of the two adjacent cells.

■ ■

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

5.1 (page 107) 1 Evaluate the importance of membranes to the homeostasis of the cell, emphasizing their various functions. The plasma membrane physically separates the interior of the cell from the extracellular environment, receives information about changes in the environment, regulates the passage of materials into and out of the cell, and communicates with other cells. ■ Biological membranes form compartments within eukaryotic cells that allow a variety of separate functions. Membranes participate in and serve as surfaces for biochemical reactions. 2 Describe the fluid mosaic model of cell membrane structure. ■ According to the fluid mosaic model, membranes consist of a fluid phospholipid bilayer in which a variety of proteins are embedded. The phospholipid molecules are amphipathic: they have hydrophobic and hydrophilic regions. The hydrophilic heads of the phospholipids are at the two surfaces of the bilayer, and their hydrophobic fatty acid chains are in the interior. ■

3 Relate properties of the lipid bilayer to properties and functions of cell membranes. In almost all biological membranes, the lipids of the bilayer are in a fluid or liquid-crystalline state, which allows the lipid molecules to move rapidly in the plane of the membrane. Proteins also move within the membrane. ■ Lipid bilayers are flexible and self-sealing and can fuse with other membranes. These properties allow the cell to transport materials

from one region of the cell to another; materials are transported in vesicles that bud from one cell membrane and then fuse with some other membrane. 4 Describe the ways that membrane proteins associate with the lipid bilayer. ■ Integral membrane proteins are embedded in the bilayer with their hydrophilic surfaces exposed to the aqueous environment and their hydrophobic surfaces in contact with the hydrophobic interior of the bilayer. Transmembrane proteins are integral proteins that extend completely through the membrane. Peripheral membrane proteins are associated with the surface of ■ the bilayer, usually bound to exposed regions of integral proteins, and are easily removed without disrupting the structure of the membrane.

5.2 (page 114) 5 Summarize the functions of membrane proteins. ■

Membrane proteins anchor cells, transport materials, act as enzymes or receptors, recognize cells and communicate with them, and structurally link cells.

■

5.3 (page 115) 6 Describe the importance of selectively permeable membranes and compare the functions of carrier proteins and channel proteins. ■ Biological membranes are selectively permeable membranes: they allow the passage of some substances but not others. By regulating

■

■

Plant cells withstand high internal hydrostatic pressure because their cell walls prevent them from expanding and bursting. Water moves into plant cells by osmosis and fills the central vacuoles. The cells swell, building up turgor pressure against the supportive cell walls. Plasma membrane

Vacuole

Learn more about the plasma membrane and membrane proteins by clicking on the figures in CengageNOW.

Vacuolar membrane (tonoplast)

5.4 (page 116) 7 Contrast simple diffusion with facilitated diffusion.

■

Diffusion is the net movement of a substance down its concentration gradient from a region of greater concentration to one of lower concentration. Diffusion and osmosis are physical processes that do not require the cell to directly expend metabolic energy. In simple diffusion through a biological membrane, solute molecules or ions move directly through the membrane down their concentration gradient. Facilitated diffusion uses specific transport proteins to move solutes across a membrane. As in simple diffusion, net movement is always from a region of higher to a region of lower solute concentration. Facilitated diffusion cannot work against a concentration gradient. Outside cell

K+ K+

K+

K+

5.5 (page 121) 9 Describe active transport, including cotransport. ■

In active transport, the cell expends metabolic energy to move ions or molecules across a membrane against a concentration gradient. For example, the sodium–potassium pump uses ATP to pump sodium ions out of the cell and potassium ions into the cell.

Higher

K+ K+

Cytoplasm

Na+

K+

Na+

K+ Cytosol

Lower

Outside cell Active transport channel

ADP + P

ATP

8 Define osmosis and solve simple problems involving osmosis; for example, predict whether cells will swell or shrink under various osmotic conditions. ■ Osmosis is a kind of diffusion in which molecules of water pass through a selectively permeable membrane from a region where water has a higher effective concentration to a region where its effective concentration is lower. ■ The concentration of dissolved substances (solutes) in a solution determines its osmotic pressure. Cells regulate their internal osmotic pressures to prevent shrinking or bursting. An isotonic solution has an equal solute concentration compared ■ to that of another fluid, for example, the fluid within the cell. ■ When placed in a hypertonic solution, one that has a greater solute concentration than that of the cell, a cell loses water to its surroundings; plant cells undergo plasmolysis, a process in which the plasma membrane separates from the cell wall. ■ When cells are placed in a hypotonic solution, one that has a lower solute concentration than the solute concentration of the cell, water enters the cells and causes them to swell.

Na+

Sodium concentration gradient

■

Potassium concentration gradient

■

passage of molecules that enter and leave the cell and its compartments, the cell controls its volume and the internal composition of ions and molecules. Membrane transport proteins facilitate the passage of certain ions and molecules through biological membranes. Carrier proteins are transport proteins that undergo a series of conformational changes as they bind and transport a specific solute. ABC transporters are carrier proteins that use energy from ATP to transport solutes. Channel proteins are transport proteins that form passageways through which water and certain ions travel through the membrane. Porins are channel proteins that form relatively large pores through the membrane for passage of water and certain solutes.

K+

Lower ■

Cytosol

K+

Higher

In cotransport, also called indirect active transport, two solutes are transported at the same time. An ATP-powered pump maintains a concentration gradient. Then a carrier protein cotransports two solutes. It transports one solute down its concentration gradient and uses the energy released to move another solute against its concentration gradient.

5.6 (page 123) 10 Compare exocytotic and endocytotic transport mechanisms. ■

The cell expends metabolic energy to carry on exocytosis and endocytosis. In exocytosis, the cell ejects waste products or secretes substances such as mucus by fusion of vesicles with the plasma membrane. This process increases the surface area of the plasma membrane.

■

■

■

■

In endocytosis, materials such as food particles are moved into the cell. A portion of the plasma membrane envelops the material, enclosing it in a vesicle or vacuole that is then released inside the cell. This process decreases the surface area of the plasma membrane. Three types of endocytosis are phagocytosis, pinocytosis, and receptor-mediated endocytosis. In phagocytosis, the plasma membrane encloses a large particle such as a bacterium, forms a vacuole around it, and moves it into the cell. In pinocytosis, the cell takes in dissolved materials by forming tiny vesicles around droplets of fluid trapped by folds of the plasma membrane. In receptor-mediated endocytosis, specific receptors in coated pits along the plasma membrane bind ligand molecules. These pits, coated by the protein clathrin, form coated vesicles by endocytosis. The vesicles fuse with lysosomes, and their contents are digested and released into the cytosol. Learn more about membrane transport by clicking on the figures in CengageNOW.

5.7 (page 127) 11 Compare the structures and functions of anchoring junctions, tight junctions, gap junctions, and plasmodesmata. ■ Cells in close contact with one another may form intercellular junctions. Anchoring junctions include desmosomes and adhering junctions; they are found between cells that form a sheet of tissue. Desmosomes spot-weld adjacent animal cells together. Adhering junctions are formed by cadherins, transmembrane proteins that cement cells together. ■ Tight junctions seal membranes of adjacent animal cells together, preventing substances from moving through the spaces between the cells. ■ Gap junctions, composed of the protein connexin, form channels that allow communication between the cytoplasm of adjacent animal cells. ■ Plasmodesmata are channels connecting adjacent plant cells. Openings in the cell walls allow the plasma membranes and cytosol to be continuous; certain molecules and ions can pass from cell to cell.

T E S T YO U R U N D E R S TA N D I N G 1. According to the fluid mosaic model, membranes consist of (a) a lipid–protein sandwich (b) mainly phospholipids with scattered nucleic acids (c) a fluid phospholipid bilayer in which proteins are embedded (d) a fluid phospholipid bilayer in which carbohydrates are embedded (e) a protein bilayer that behaves as a liquid crystal 2. Transmembrane proteins (a) are peripheral proteins (b) are receptor proteins (c) extend completely through the membrane (d) extend along the surface of the membrane (e) are secreted from the cell 3. Which of the following is not a function of the plasma membrane? (a) transports materials (b) helps structurally link cells (c) has receptors that relay signals (d) anchors the cell to the extracellular matrix (e) manufactures proteins 4. ABC transporters (a) use the energy of ATP hydrolysis to transport certain ions and sugars (b) are important in facilitated diffusion of certain ions (c) are a small group of channel proteins (d) are found mainly in plant cell membranes (e) permit passive diffusion through their channels 5. When plant cells are in a hypotonic medium, they (a) undergo plasmolysis (b) build up turgor pressure (c) wilt (d) decrease pinocytosis (e) lose water to the environment 6. A laboratory technician accidentally places red blood cells in a hypertonic solution. What happens? (a) they undergo plasmolysis (b) they build up turgor pressure (c) they swell (d) they pump solutes out (e) they become dehydrated and shrunken

7. Which of the following processes requires the cell to expend metabolic energy directly (for example, from ATP)? (a) osmosis (b) facilitated diffusion (c) all forms of carrier-mediated transport (d) active transport (e) simple diffusion 8. Electrochemical gradients (a) power simple diffusion (b) are established by pinocytosis (c) are necessary for transport by aquaporins (d) are established by concentration gradients (e) are a result of both an electric charge difference and a concentration difference between the two sides of the membrane 9. In cotransport (indirect active transport) (a) a uniporter moves a solute across a membrane against its concentration gradient (b) the movement of one solute down its concentration gradient provides energy for transport of some other solute up its concentration gradient (c) a channel protein moves ions by facilitated diffusion (d) osmosis powers the movement of ions against their concentration gradient (e) sodium is directly transported in one direction, and potassium is indirectly transported in the same direction 10. A cell takes in dissolved materials by forming tiny vesicles around fluid droplets trapped by folds of the plasma membrane. This process is (a) indirect active transport (b) pinocytosis (c) receptormediated endocytosis (d) exocytosis (e) facilitated diffusion 11. Anchoring junctions that hold cells together at one point as a spot weld does are (a) tight junctions (b) adhering junctions (c) desmosomes (d) gap junctions (e) plasmodesmata

12. Junctions that permit the transfer of water, ions, and molecules between adjacent plant cells are (a) tight junctions (b) adhering junctions (c) desmosomes (d) gap junctions (e) plasmodesmata 13. Label the cell membrane. Consult Figure 5-6 to check your answers.

CRITICAL THINKING 1. Why can’t larger polar molecules and ions diffuse through the plasma membrane? Would it be advantageous to the cell if they could? Explain. 2. Describe one way that an ion gradient can be established and maintained. 3. Most adjacent plant cells are connected by plasmodesmata, whereas only certain adjacent animal cells are associated through gap junctions. What might account for this difference? 4. EVOLUTION LINK. Hypothesis: the evolution of biological membranes was an essential step in the origin of life. Give arguments supporting (or challenging) this hypothesis. 5. EVOLUTION LINK. Transport proteins have been found in all biological membranes. What hypothesis could you make regarding whether these molecules evolved early or later in the history of cells? Argue in support of your hypothesis.

6. SCIENCE, TECHNOLOGY, AND SOCIETY. Biologists know quite a bit about the components that make up plant cell walls, but they do not have enough data to model the components of the cell wall as a system. Would you be in favor of investing government funds in the research and technology that would allow researchers to model the cell wall? Why or why not? Would your answer be different if you considered that a better understanding of the cell wall might allow biologists to manipulate plant growth, ripening of fruit, and the texture of certain foods? Additional questions are available in CengageNOW at www.cengage.com/ login.

Cell Communication

From I. Joint et al., Science 298:1207

6

Cell-to-cell communication across the prokaryote– eukaryote boundary. Spores of the green seaweed Enteromorpha (Domain Eukarya) attach to biofilmforming bacteria (Domain Bacteria) in response to chemical compounds released by the bacteria. The bacteria (blue) were stained and visualized with blue light. The spores appear red because of the fluorescence of chlorophyll within them.

KEY CONCEPTS

T

6.1 Cells communicate by signaling one another, a com-

nisms for transmitting information between cells, tissues, and organs

plex process that involves production of signaling molecules, reception of the signal, signal transduction, and a response.

have evolved, including electrical signaling and many types of chemical

6.2 Cells signal one another using chemical compounds such as neurotransmitters, hormones, and other signaling molecules.

o maintain homeostasis, the cells of a multicellular organism must continuously communicate with one another. Many different mecha-

signaling. Organisms also communicate with other members of their species by secreting chemical signals. For example, bacteria release chemical signals that diffuse among nearby bacteria. As the population of bacteria increases, the concentration of the chemical signal increases. Through a

6.3 A signaling molecule binds to a receptor molecule on

process known as quorum sensing, the bacteria sense when a certain criti-

the cell surface or inside the target cell.

cal concentration of a signal molecule is reached. The bacteria respond by

6.4 In signal transduction, a cell converts an extracellular

activating a specific biological process. For example, they may form a bio-

signal into an intracellular signal and relays the signal, leading to some change in the cell (the response).

film, a community of microorganisms attached to a solid surface. Forming

6.5 The cell responds to signals by opening or closing ion channels; altering enzyme activity, which leads to metabolic changes and other alterations in cell activity; and by activating or inhibiting specific genes.

6.6 Similarities in cell communication among diverse organisms suggest that the molecules and mechanisms used in information transfer evolved long ago.

a biofilm requires the coordinated activity of numerous bacteria. Over billions of years, elaborate systems of cell signaling have evolved. Organisms of different species, and even different kingdoms and domains, communicate with one another. For example, the chemical signals released by bacteria can be intercepted by other organisms. During its life cycle, the green seaweed Enteromorpha produces spores that move about and temporarily attach to a surface. The spores sense a chemical signal released by bacteria that form biofilms. In response to the chemical

signal, the spores move toward the biofilm and attach to individual bacteria that are part of the biofilm surface (see opening photograph). Thousands of chemical reactions are involved in responding to signal molecules and regulating the communication among molecules necessary to maintain homeostasis. Many researchers are working on the molecular level to understand how proteins receive messages and relay signals. They are learning how proteins act as molecular switches, activating and deactivating molecules in complex signaling pathways.

communication. Biologists, biochemists, physicists, and scientists from many other disciplines are working together to understand how elaborate signaling systems within the cell interact to maintain homeostasis. They are studying how information is transferred between cells at the tissue and organ levels, and throughout the organism. Faulty signaling in cells and between cells can cause a variety of diseases, including cancer and diabetes. In Chapter 1, we introduced three basic themes of biology. One of these, transmission of information, is the main focus of this chapter. In this chapter, we discuss how cells send and receive sig-

16.6 mm

understand the intricate, dynamic interactions involved in cell

G. Gerisch et al./Max-Planck Institute

Some cell biologists are using a systems biology approach to

FIGURE 6-1 Cell signaling in cellular slime molds When food is scarce, the amoeba-like cellular slime mold Dictyostelium secretes the chemical compound cyclic AMP (cAMP). The slime molds respond to this chemical signal by aggregating. Converging streams of hundreds of individuals come together and form a multicellular colony.

nals. We consider how information crosses the plasma membrane and is transmitted through signaling systems. We describe some of the responses that cells make. Finally, we discuss the evolution of cell communication. Increased understanding of the mechanisms of cell communication may suggest new strategies for preventing and treating diseases.

6.1 CELL COMMUNICATION: AN OVERVIEW ■ ■ LEARNING OBJECTIVE 1

Describe the four main processes essential for cells to communicate.

When food is scarce, the amoeba-like cellular slime mold Dictyostelium secretes the compound cyclic adenosine monophosphate (cAMP). This chemical compound diffuses through the cell’s environment and binds to receptors on the surfaces of nearby cells. The activated receptors send signals into the cells that result in movement toward the cAMP. Hundreds of slime molds come together and form a multicellular slug-shaped colony (FIG. 6-1; also see Fig. 26-20). (When conditions are favorable, the cells of the slug form a stalked fruiting body with spores at the top; when released, each spore gives rise to an amoeba-like cell.) Even though they do not move from one place to another, plants also communicate with each other. For example, diseased maple trees send airborne chemical signals that are received by uninfected trees nearby. Cells of the uninfected trees respond by increasing their chemical defenses so that they are more resistant to the disease-causing organisms. Plants also send signals to insects. When tobacco plants are attacked by herbivorous insects, the

plants release volatile chemicals. In response to these signals, the insects lay fewer eggs, thus reducing the number of insects feeding on the plants. Predator insects that eat the eggs of the herbivorous insects respond to the plant signals by eating more of the herbivorous insect eggs. Thus, natural selection has resulted in plant signals that herbivorous insects detect and avoid and that carnivorous insects detect and approach. This system of information transfer helps protect plants from herbivorous insects. To survive, organisms must receive signals from the outside environment and effectively respond to them. In order to grow, develop, and function, the cells of a multicellular organism must also communicate with one another. In plants and animals, hormones and other regulatory molecules serve as important chemical signals between various cells and organs. In animals, neurons (nerve cells) transmit information electrically and chemically. The term cell signaling refers to the mechanisms by which cells communicate with one another. If the cells are physically close to one another, a signaling molecule on one cell may combine with a receptor (a macromolecule that binds with signaling molecules) on another cell. Most commonly, cells communicate by sending chemical signals over some distance. As we will discuss, cell signaling must be precisely regulated. Cell signaling involves a sequence of four main processes (FIG. 6-2). We summarize them here and discuss them in more detail in the following sections of this chapter. 1. A cell must send a signal. In chemical signaling, a cell must synthesize and release signaling molecules. For example, specialized cells in the vertebrate pancreas secrete the hormone insulin. If the target cells, the cells that can respond to the signal, are not in close proximity, the signal must be transported to them. The circulatory system transports insulin to target cells

throughout the body. Next, target KEY POINT When a signaling molecule binds to a receptor molecule on a target cell, the cells must receive, relay, and respond receptor activates a signal transduction pathway, leading to some response to the information signaled. in the cell. 2. Reception. Reception is the process of receiving an incoming signal. Receptors are large proteins or glycoproteins that bind with signaling molecules. Many types of signaling 1 Cell sends signal molecules do not actually enter the target cell. They bind to specific reSignaling ceptors on the surface of the target molecules cell. Insulin, for example, binds to Receptor insulin receptors, which are transmembrane receptors. 3. Signal transduction is the process 2 Reception by which a cell converts an extracellular signal into an intracellular Signaling signal and relays the signal, leading molecule to a cellular response. Signal transA duction typically involves a chain Signaling of molecules that relay information. molecule When insulin binds to an insulin reB Target ceptor, the signal is relayed through cell several different signaling pathways. 3 Signal Signaling transduction 4. Response. The final molecule in the molecule signaling pathway converts the sigC nal into a response that alters some cell process. For example, as we will discuss in Chapter 49, insulin stimulates cells to take up glucose from the Protein Protein that Enzyme blood. This response lowers the conregulates a gene centration of glucose in the blood. Insulin also helps regulate fat and protein metabolism. Many signaling 4 Response Altered Change in some Specific gene molecules stimulate ion channels in membrane metabolic process activated or repressed permeability the plasma membrane to open or close. Still other signaling molecules activate or inhibit specific genes in FIGURE 6-2 Animated Overview of cell signaling the nucleus. These responses can result in changes in cell division and other aspects of cell development. Cells communicate in several ways, including directly through cell After a signaling molecule has done its job, its action must be junctions, by way of electrical signals, temporary cell-to-cell constopped, so during the signaling process, certain mechanisms tact, and chemical signals. Recall from Chapter 5 that gap junctions operate to terminate the signal. in animal cells allow rapid chemical and electrical communication between adjacent cells. For example, the gap junctions between Review cardiac muscle cells of the heart wall allow the rapid flow of ions ■ What is the sequence of events that takes place in cell signaling? necessary for synchronized contraction. Plasmodesmata between Describe each process. adjacent plant cells also allow signal molecules to pass quickly from one cell to another. Cells that are not directly connected also communicate with 6.2 SENDING SIGNALS one another. In animals, some neurons communicate with electrical signals. Most neurons, however, signal one another by releasing ■ ■ LEARNING OBJECTIVE chemical compounds called neurotransmitters (FIG. 6-3a). Neurotransmitter molecules diffuse across synapses, tiny gaps between 2 Compare three types of signaling molecules: neurotransmitters, hormones, and local regulators. neurons. More than 60 different neurotransmitters have been

hormones into the surrounding interstitial fluid. Typically, hormones Receptor diffuse into capillaries and are transSignaling ported by the blood to target cells molecules (FIG. 6-3c). Target Some cells produce local regulaneuron tors that signal cells in close proximSignaling ity. A local regulator is a signaling neuron molecule that diffuses through the interstitial fluid, the fluid surrounding the cells, and acts on nearby cells. (b) Some cells signal one another Receptor by making direct contact. This is called paracrine regulation (FIG. 6-3d). Some local regulators are considered hormones. Local regulaNucleus tors include local chemical mediators such as growth factors, histamine, nitric oxide, and prostaglandins. More Signaling Target than 50 growth factors, typically cell cell peptides, stimulate cell division and normal development in specific types Receptor Signaling (c) Many hormones are transported of cells. molecules by the blood to target cells. (hormone) Histamine is a local regulator that is stored in certain cells of the immune system and is released in response to allergic reactions, injury, or infection. Histamine causes blood Hormone transported vessels to dilate and capillaries to beEndocrine Target in blood come more permeable. Nitric oxide cell cell (NO), another local regulator, is a gas produced by many types of cells, (d) In paracrine regulation, a local Signaling Receptor molecules regulator diffuses to target cells. including plant and animal cells. (local Nitric oxide released by cells lining regulator) blood vessels relaxes smooth muscle in the blood vessel walls. As a result, the blood vessels dilate, decreasing blood pressure. Signaling Prostaglandins are local horTarget cell cell mones that are paracrine regulators. Prostaglandins modify cAMP levels and interact with other signaling molecules to regulate metabolic FIGURE 6-3 Some types of cell signaling activities. For example, some prostaglandins stimulate smooth Different types of cells may communicate in different ways. muscle to contract. (a) Neurons transmit signals across synapses.

identified, including acetylcholine, norepinephrine, dopamine, serotonin, and several amino acids and peptides. Cells synthesize many different types of chemical signals and deliver them in various ways. In animals, certain cells in the immune system produce specific chemical compounds that are displayed on the cell surface. These cells recognize the chemical signals and communicate with one another by making direct contact (FIG. 6-3b). Specialized cells in plants and animals produce signaling molecules called hormones. Hormones may be synthesized by neighboring cells or by specialized organs or tissues some distance from the target cells. In animals, many hormones are produced by endocrine glands. These glands have no ducts; they secrete their

Review ■ ■ ■

What are neurotransmitters? How are hormones typically transported to target cells? What is paracrine regulation?

6.3 RECEPTION ■ ■ LEARNING OBJECTIVES 3 Identify mechanisms that make reception a highly specific process. 4 Briefly compare ion channel–linked receptors, G protein–linked receptors, enzyme-linked receptors, and intracellular receptors.

Hundreds of different types of signaling molecules are present in the interstitial fluid, the tissue fluid that surrounds the Receptor cells of a multicellular organism. How do cells know which messages are for them? Signaling molecules The answer is that each type of cell is genetically programmed to receive and respond to specific types of signals. Which signals a cell responds to depends on the specific receptors it is programmed to synthesize. A signaling molecule, such as insulin, that binds to a specific receptor acts as a liNucleus gand. A ligand is a molecule, other than an enzyme, that binds specifically to a macromolecule (usually a protein), forming a macromolecule-ligand complex. The com(a) Hydrophilic signaling molecules bind to receptors in the plasma membrane. plex triggers a biological response. Most ligands are hydrophilic molecules that bind to protein receptors on the surface of tarSignaling get cells (FIG. 6-4a). molecules Some signaling molecules are small Receptor enough or sufficiently hydrophobic to move through the plasma membrane and enter the cell (FIG. 6-4b). These signaling molecules bind with intracellular receptors. Reception occurs when a signaling molecule binds to a specific receptor protein on the surface of, or inside, a target cell. The signaling molecule activates the receptor. A receptor on the cell surface generally has at least three domains. Recall from Chapter 3 that in biochemistry, the term domain refers to a structural and functional (b) Hydrophobic signaling molecules cross the plasma membrane and bind region of a protein. The external domain with receptors inside the cell. is a docking site for a signaling molecule. A second domain extends through the FIGURE 6-4 Cell-surface and intracellular receptors plasma membrane, and a third domain is a Hydrophilic (water-soluble) signaling molecules cannot pass through the plasma membrane. They bind to receptors on the cell surface. Hydrophobic (lipid-soluble) signaling molecules pass through the “tail” that extends into the cytoplasm. The plasma membrane and bind with receptors in the cytosol or nucleus. tail transmits the signal to a molecule inside the cell. Reception is highly selective. Each ment proteins that are activated by red light. Activation can lead type of receptor has a specific shape. The receptor binding site is to changes, such as flowering. Plants, some algae, and at least some somewhat like a lock, and signaling molecules are like different animals have cryptochromes, pigments that absorb blue light. keys. Only the signaling molecule that fits the specific receptor can Cryptochromes play a role in biological rhythms. influence the metabolic machinery of the cell. Receptors are important in determining the specificity of cell communication. Different types of cells can produce different types of recepCells regulate reception tors. Any one cell makes many different receptors. Furthermore, a An important mechanism that cells use to regulate reception is incell may synthesize different kinds of receptors at different stages creasing or decreasing the number of each type of receptor. Dependin its life cycle or in response to different conditions. Another coning on the needs of the cell, receptors are synthesized or degraded. sideration is that the same signal can have different meanings for For example, when the concentration of the hormone insulin is too various target cells. high for an extended period, cells decrease the number of their inSome receptors are specialized to respond to signals other sulin receptors. This process is called receptor down-regulation. than chemical signals. For example, in the vertebrate eye, a recepIn the case of insulin, receptor down-regulation suppresses the tor called rhodopsin is activated by light. Rhodopsin is part of a sensitivity of target cells to the hormone. Insulin stimulates cells to signal transduction pathway that leads to vision in dim light. Plants take in glucose by facilitated diffusion, so receptor down-regulation and some algae have phytochromes, a family of blue-green pig-

decreases the ability of cells to take in glucose. Receptor downregulation often involves transporting receptors to lysosomes, where they are destroyed. Receptor up-regulation occurs in response to low hormone concentrations. A greater number of receptors are synthesized, and their increased numbers on the plasma membrane make it more likely that the signal will be received by a receptor on the cell. Receptor up-regulation thus amplifies the signaling molecule’s effect on the cell. Receptor up-regulation and down-regulation are controlled in part by signals to genes that code for the receptors.

Three types of receptors occur on the cell surface Three main types of receptors on the cell surface are ion channel– linked receptors, G protein–linked receptors, and enzyme-linked receptors. Ion channel–linked receptors convert chemical signals into electrical signals Ion channel–linked receptors are found in the plasma membrane. These receptors, which have been extensively studied in neurons and muscle cells, convert chemical signals into electrical signals (FIG. 6-5a). The receptor itself serves as a channel. Ion channel–linked receptors are also called ligand-gated channels, which means that the ion channel opens or closes in response to the binding of the signaling molecule (ligand). Part of the receptor (protein) that makes up the channel forms the gate. The receptor responds to certain signals by changing its shape, opening or closing the gate. Typically, the gate of an ion channel remains closed until a ligand binds to the receptor. Neurotransmitters released by neurons in the nervous system are signaling molecules. For example, acetylcholine is a neurotransmitter that binds to an acetylcholine receptor. This receptor is a ligand-gated sodium ion channel that is important in muscle contraction. When acetylcholine binds to the receptor, the channel opens, allowing sodium ions to enter the cell. The influx of sodium ions decreases the electric charge difference across the membrane (depolarization), which can lead to muscle contraction. After a brief time, the ligand dissociates from the receptor and the gate closes the channel. As we will discuss in Chapter 41, some ion channels, called voltage-activated channels, are regulated by electrical signals. G protein–linked receptors link signaling molecules to signal transduction pathways G protein–linked receptors (also called G protein–coupled receptors) are a large family of transmembrane proteins that loop back and forth through the plasma membrane seven times (FIG. 6-5b). The receptor consists of seven transmembrane alpha helices connected by loops that extend into the cytosol or outside the cell. G protein–linked receptors couple certain signaling molecules to various signal transduction pathways inside the cell. The outer part of the receptor has a binding site for a signaling molecule, and the part of the receptor that extends into the cytosol has a binding site for a specific G protein.

G proteins bind guanine nucleotides. When a signaling molecule binds with a G protein–linked receptor, the receptor changes shape. This change allows the G protein to associate with the receptor. G protein–linked receptors are found in all eukaryotes. These receptors bind with hundreds of different signaling molecules. Some of the many critical processes that depend on G protein–linked receptors are vision, sense of smell, regulation of mood and behavior, and regulation of the immune system. As you might imagine, understanding how G protein–linked receptors work is medically important. About 60% of prescription medications currently in use act on these receptors. More than 900 G protein–linked receptors have been identified in mammals, and more than 400 of these receptors are potential targets for pharmaceutical interventions. Enzyme-linked receptors function directly as enzymes or are linked to enzymes Enzyme-linked receptors are transmembrane proteins with a binding site for a signaling molecule outside the cell and an enzyme component inside the cell. Recall from Chapter 3 that enzymes catalyze specific chemical reactions; most enzymes are proteins. Some enzyme-linked receptors do not have an enzyme component but have a binding site for an enzyme. Several groups of enzyme-linked receptors have been identified. Proteins called tyrosine kinases make up a major group of enzyme-linked receptors. A kinase is an enzyme that transfers the terminal phosphate group from ATP to a substrate (the specific substance on which an enzyme acts). This process is called phosphorylation. A tyrosine kinase is an enzyme that catalyzes the transfer of phosphate groups from ATP to a specific tyrosine that is part of a protein. (Tyrosine is an amino acid.) The tyrosine kinase enzyme part of the receptor is the domain that extends into the cytosol. Tyrosine kinase receptors bind certain hormones, including insulin and growth factors, such as nerve growth factor. These receptors regulate many cellular processes and are important in development. Before the tyrosine kinase can phosphorylate a signaling protein in the cell, it must first be activated. When signal molecules bind to two tyrosine kinase receptors, the receptors move closer together in the plasma membrane and pair, forming a dimer. A conformational change (change in shape) takes place, which allows the tyrosine kinase part of each receptor to add a phosphate from an ATP molecule to certain tyrosines on the other member of the dimer (FIG. 6-5c). Once activated, tyrosine kinase enzymes can phosphorylate signaling proteins inside the cell (discussed in the next section). Many plant cell-surface receptors are enzyme-linked receptors. Brassinosteroids (BRs), a group of plant steroid hormones, regulate many plant processes, including cell division, cell elongation, and flower development (discussed in Chapter 38). Unlike animal steroid hormones, which typically bind with intracellular receptors, BRs bind with a protein kinase receptor in the plasma membrane. In plant protein kinase receptors, serine and threonine, rather than tyrosine, appear to be the amino acids that are phosphorylated. The gas ethylene is a plant hormone that regulates a variety of processes, including seed germination and ripening of fruit. Ethylene is also important in plant responses to stressors. The ethylene receptor has two components. Each component has a domain that

Signaling molecule

Extracellular fluid

Na+

Receptor

Cytosol

1

Ion channel is closed.

2

When signaling molecule binds to receptor, the channel opens. Sodium ions enter cell.

(a) Ion channel–linked receptor

Signaling molecule

Receptor G protein

Enzyme

Cytosol

GTP

GDP 1

In inactive state, the three subunits of G protein are joined.

2

When signaling molecule (ligand) binds to receptor, the ligand–receptor complex associates with G protein. This causes GDP to be replaced with GTP. One subunit of G protein then separates from the other two parts.

(b) G protein–linked receptor

Dimer

Signaling molecule Receptor

Enzyme domain of receptor

Tyr

Tyr

Tyr

Tyr

Tyr

Tyr

Tyr

Tyr

Cytosol 8 ATP

8 ADP

Protein kinase sites (inactive) 1

Receptors are in inactive state.

(c) Enzyme-linked receptors

FIGURE 6-5 Three types of cell-surface receptors

P

Tyr

Tyr

P

P

Tyr

Tyr

P

P

Tyr

Tyr

P

P

Tyr

Tyr

P

Protein kinase sites (active) 2

When signaling molecule (ligand) binds to receptors, receptors are enzymatically phosphorylated. Phosphate comes from ATP.

is an enzyme, a histidine kinase, that extends into the cell. Receptors with histidine kinase domains are also present in bacterial and yeast cells.

Some receptors are located inside the cell Certain receptors are found in the cytosol or in the nucleus. Most of these intracellular receptors are transcription factors, proteins that regulate the expression of specific genes. The signaling molecules that bind with intracellular receptors are small, hydrophobic molecules that can diffuse across the membranes of target cells (see Fig. 6-4b). In animal cells, most steroid hormones, such as the molting hormone ecdysone in insects and cortisol in vertebrates, enter target cells and combine with receptor molecules in the cytosol. Vitamins A and D and nitric oxide also bind with intracellular receptors. After binding, the ligand–receptor complex moves into the nucleus. Thyroid hormones (which are not steroids) bind to receptors already bound to DNA inside the nucleus.

Review ■ ■

■

How does a receptor “know” which signaling molecules to bind? Under what conditions might receptor up-regulation occur? receptor down-regulation? What are the three main types of cell-surface receptors? (Briefly describe each.)

6.4 SIGNAL TRANSDUCTION ■ ■ LEARNING OBJECTIVES 5 6

Compare the actions of the main types of receptors in signal transduction. Trace the sequence of events in signal transduction for each of the following second messengers: cyclic AMP, inositol trisphosphate, diacylglycerol, and calcium ions.

As we have discussed, many regulatory molecules transmit information to the cell’s interior without physically crossing the plasma membrane. Instead, they activate membrane proteins, which then transduce the signal. The first component in a signal transduction pathway is typically the receptor, which may be a transmembrane protein with a domain exposed on the extracellular surface. Each type of receptor activates a different signal transduction pathway. In a typical signaling pathway, a signaling molecule binds with a cell-surface receptor and activates it by changing the shape of the receptor tail that extends into the cytoplasm. The signal may then be relayed through a sequence of proteins that are intracellular signaling molecules. Typically, the proteins in this chain are protein kinases. The chain of signaling molecules in the cell that relays a signal is called a signaling pathway or signaling cascade. During signal transduction, the original signal is amplified. More than 3000 signaling proteins have been identified in the cells of mammals alone. Malfunctions in signal transduction pathways have been linked to major diseases, including cancer, heart disease, diabetes, and autoimmune diseases.

Signaling molecules can act as molecular switches Each component in a signaling pathway acts as a molecular “switch,” which can be in an active (“on”) state or an inactive (“off ”) state. When an intracellular signaling molecule receives a signal, it is activated (turned on). Another process then occurs that inactivates (turns off ) the signaling molecule. Every activated molecule in a signaling pathway must be inactivated in order to transmit a new signal. Molecular switches are typically regulated by the addition or removal of phosphate groups. Each time a signal molecule binds to a receptor, a signaling pathway is turned on. As the signal is transmitted to each protein kinase in the chain, the protein is phosphorylated. Recall that in phosphorylation, an enzyme transfers phosphate groups from one molecule to another. As phosphate is transferred, the signal passes from protein kinase 1 to protein kinase 2 to protein kinase 3, and on down the chain (FIG. 6-6). Phosphorylation typically activates a protein kinase, although in some cases phosphorylation inhibits protein kinase activity. A signaling pathway in which a series of protein kinase molecules are phosphorylated is referred to as a protein kinase cascade. The last protein kinase in the cascade activates the target protein by phosphorylation. The target protein alters some process in the cell. After the signal passes from protein kinase 1 to protein kinase 2, protein kinase 1 must be inactivated. Each molecule must be available to transmit a new signal. A phosphatase is an enzyme that catalyzes the removal of a phosphate group by hydrolysis; the process is called dephosphorylation. Protein phosphatases help regulate protein kinase cascades. Just as a cell contains hundreds of different protein kinases, it also contains many types of protein phosphatases. Rapid removal of phosphate groups by protein phosphatases is an important regulatory mechanism for protein kinase cascades. Such regulation ensures that these pathways operate only in response to the binding of a signal molecule to a receptor.

Ion channel–linked receptors open or close channels The gates of many ion channels remain closed until a ligand binds to the receptor. For example, when the neurotransmitter acetylcholine binds to an acetylcholine receptor (which is an ion channel–linked receptor), the channel opens, allowing sodium ions to enter the cell (see Fig. 6-5a). Depending on the type of cell, the influx of sodium ions can result in transmission of a neural impulse or in muscle contraction. Gamma-aminobutyric acid (GABA) is a neurotransmitter that binds to GABA receptors. One class of GABA receptors consists of ligand-gated chloride ion channels. When GABA binds to the receptor, the channel opens. Chloride ions, which are negatively charged, rush out of or into the neuron, depending on the conditions (electrochemical gradient) in the cell. Typically, chloride ions leave the cell, which inhibits transmission of neural impulses. Thus, GABA inhibits neural signaling. Barbiturates and benzodiazepine drugs such as Valium bind to GABA receptors. When this occurs, lower amounts of GABA are required to open the chloride channels and inhibit neural impulses. This action results in a tranquilizing effect. (Not all GABA receptors are themselves ion

KEY POINT

Many signaling pathways are protein kinase cascades in which each protein kinase in the pathway is activated by the addition of a phosphate group.

Signaling molecule

Receptor

P

Tyr

Tyr

P

P

Tyr

Tyr

P

P

Tyr

Tyr

P

P

Tyr

Tyr

P

1

Signaling molecules bind with receptors, activating them.

2

Active protein kinase 1

Inactive protein kinase 1

Receptor activates protein kinase 1 by phosphorylation.

P

Inactive protein kinase 2

ATP

3

Active protein kinase 2

ADP

Active protein kinase 1 then activates protein kinase 2 by phosphorylation.

P

Inactive protein kinase 3

ATP

4

Active protein kinase 3

ADP

Active protein kinase 2 activates protein kinase 3 by phosphorylation.

P 5

Inactive protein

ATP

ADP

Active protein

Active protein kinase 3 phosphorylates a specific protein that alters some cell process.

P

Alters some cell process

FIGURE 6-6 A phosphorylation cascade When the receptor is activated, each protein kinase in a signaling pathway activates the next protein kinase in the pathway by phosphorylation. The phosphate group comes from ATP. Addition of a phosphate group typically changes the shape of the molecule. Activation of the

channels; some are G-protein–linked receptors or enzyme-linked receptors that trigger a series of reactions resulting in activation of other proteins that serve as ion channels.)

G protein–linked receptors initiate signal transduction As discussed in the last section, G protein–linked receptors activate G proteins, a group of regulatory proteins important in many signal transduction pathways. These proteins are found in yeasts, protists, plants, and animals. They are involved in the action of

last protein in the chain changes some cell process or turns on (or turns off) specific genes. (P represents phosphate.) Note that the number of protein kinases varies from pathway to pathway.

some plant and many animal hormones. Some G proteins regulate channels in the plasma membrane, allowing ions to enter or exit the cell. Other G proteins are involved in the perception of sight and smell. In 1994, Alfred G. Gilman of the University of Texas and Martin Rodbell of the National Institute of Environmental Health Sciences were awarded the Nobel Prize in Physiology or Medicine for their groundbreaking research on G proteins. In its inactive state, the G protein consists of three subunits that are joined (see Fig. 6-5b). One subunit is linked to a molecule of guanosine diphosphate (GDP), a molecule similar to ADP but containing the base guanine instead of adenine. When a sig-

naling molecule binds with the receptor, the GDP is released and is replaced by guanosine triphosphate (GTP), a nucleotide that, like ATP, functions in energy transactions. The subunit of the G protein that is linked to the GTP separates from the other two subunits. The G protein subunit linked to the GTP is a GTPase, an enzyme that catalyzes the hydrolysis of GTP to GDP. This action, a process that releases energy, deactivates the G protein. Now in its inactive state, the G protein subunit rejoins the other two subunits. When activated, a G protein initiates signal transduction by binding with a specific protein in the cell. In some cases, G proteins directly activate enzymes that catalyze changes in certain proteins. These changes lead to alterations in cell function. More commonly, the signaling molecule serves as the first messenger, and information is relayed by way of the G protein to a second messenger.

Second messengers are intracellular signaling agents Second messengers are ions or small molecules that relay signals inside the cell. More than 15 second messengers have been identified in mammals. When receptors are activated, second messengers are produced in large quantities. Second messengers rapidly diffuse through the cell (or membrane), relaying the signal. Thus, second messengers amplify the signal. Second messengers pass the signal along to other signaling proteins or target proteins. The signal typically passes through a

KEY POINT

chain of proteins and other molecules. The last molecule in the sequence activates the final response. Second messengers are not enzymes, but some regulate specific enzymes, such as protein kinases. Others bind to ion channels, opening or closing them. Cyclic AMP is a second messenger Most G proteins shuttle a signal between the receptor and a second messenger. In many signaling cascades in prokaryotic and animal cells, the second messenger is cyclic AMP (cAMP) (FIG. 6-7). Researcher Earl Sutherland identified cAMP as a second messenger in the 1960s and was awarded the 1971 Nobel Prize in Physiology or Medicine for his pioneering work. When the G protein undergoes a conformational change (change in shape), it binds with and activates adenylyl cyclase, an enzyme on the cytoplasmic side of the plasma membrane. The type of G protein that activates adenylyl cyclase is known as a stimulatory G protein, or Gs. (Some G proteins, denoted as Gi, inhibit enzymes.) Note that the G protein couples the ligand–receptor complex to adenylyl cyclase action (see Fig. 6-7). When activated, adenylyl cyclase catalyzes the production of cAMP from ATP (FIG. 6-8). By coupling the signaling molecule– receptor complex to an enzyme that generates a signal, G proteins amplify the effects of signaling molecules and many secondmessenger molecules are rapidly produced. The pathway is regulated, in part, by phosphodiesterase, an enzyme that converts cAMP to adenosine monophosphate (AMP). This action is an off switch that rapidly inactivates cAMP when the receptor becomes inactive.

Cyclic AMP (cAMP), a second messenger, relays a signal from the plasma membrane to proteins in the cytosol. Extracellular fluid

Signaling molecule (first messenger) Receptor G protein 1

Signaling molecule combines with G protein–linked receptor. G protein activates adenylyl cyclase, which then catalyzes formation of cAMP from ATP.

Plasma membrane Cytosol

GTP

ATP 2

3

Signal is relayed by second messengers.

Response: Some cell process is altered.

Adenylyl cyclase

cAMP Second messenger

cAMP

cAMP

cAMP

Protein

Protein

Protein

Affects gene activity

Opens or closes ion channels

Alters metabolism

FIGURE 6-7 Signal relay by a second messenger: An overview

NH2

O –

O

O

O

P –

O

˜

O

P O

–

˜

O

O

CH2

O

P –

C

N

C

O H

OH

OH

N

CH2 O

C

C

O

H

H

O

OH

Phosphodiesterase

O

O

–

Adenylyl cyclase

Plasma membrane Cytosol

C N

GDP 1

CH

Signaling molecule binds with G protein–linked receptor in plasma membrane.

N

N

C

Signaling molecule Receptor

Adenylyl cyclase

G protein separates

N

C

C N

H2O

O P

Extracellular fluid

Signaling molecule Receptor

NH2

O

O

CH N

cAMP

–

–

N

NH2

N

P

When a signaling molecule binds to a G protein– linked receptor, a G protein is activated; cyclic AMP is produced and activates a signaling pathway, leading to a change in some cell process.

G protein

H

P

O

KEY POINT

C

ATP

Adenylyl cyclase P

N

CH2

GTP

CH N

2

Signal molecule–receptor complex activates G protein. GDP is replaced by GTP.

AMP

O

H

H

OH

OH

Adenylyl cyclase activated

Receptor

FIGURE 6-8 Synthesis and inactivation of cyclic AMP Cyclic AMP (cAMP) is a second messenger produced from ATP. The enzyme adenylyl cyclase catalyzes the reaction. Cyclic AMP is inactivated by the enzyme phosphodiesterase, which converts it to adenosine monophosphate (AMP). GTP

FIGURE 6-9 illustrates in more detail the sequence of events in a signaling pathway involving G protein and cyclic AMP. Cyclic AMP activates certain protein kinase enzymes, particularly a group referred to as protein kinase A. Recall that protein kinases add phosphate groups to target proteins. When a protein is phosphorylated, its function is altered and it triggers a chain of reactions leading to some response in the cell, such as a metabolic change. The substrates, the substances on which an enzyme acts, for protein kinases are different in various cell types. Consequently, the effect of the enzyme varies depending on the substrate. For example, in skeletal muscle cells, protein kinase A activates enzymes that break down glycogen to glucose, providing the muscle cells with energy. In certain neurons in the brain, the same enzyme acti-

cAMP ATP Protein kinase A

Protein

P Phosphorylated protein

Alters some cell process 3

G protein activates adenylyl cyclase, which then catalyzes synthesis of cAMP. cAMP activates protein kinase A, which then phosphorylates specific proteins, leading to some response in cell.

FIGURE 6-9 Signal transduction involving a G protein and cyclic AMP

vates the reward system by regulating action of the neurotransmitter dopamine. We can summarize the sequence of events beginning with the binding of the signaling molecule to the receptor and leading to a change in some cell function:

Some G proteins use phospholipids as second messengers Certain signaling molecule–receptor complexes activate a G protein that then activates the membrane-bound enzyme phospholipase C (FIG. 6-10). This enzyme splits a membrane phospholipid, PIP2 (phosphotidylinositol-4,5-bisphosphate), into two products, inositol trisphosphate (IP3) and diacylglycerol (DAG). Both act as second messengers. DAG remains in the plasma membrane, where in combination with calcium ions, it activates protein kinase C enzymes. Depending on the type of cell and the specific protein kinase C, the response of the cell can include growth, a change in cell pH,

signaling molecule (first messenger) binds to G protein–linked receptor ¡ activates G protein ¡ activates adenylyl cyclase ¡ catalyzes the formation of cAMP (second messenger) ¡ activates protein kinase ¡ phosphorylates proteins ¡ response in cell

KEY POINT

When activated by a G protein, phospholipase C splits the phospholipid PIP2, producing the second messengers DAG and IP3. 1

Signaling molecule binds with G protein–linked receptor and activates a G protein.

2

Signal molecule–receptor complex activates phospholipase C. Extracellular fluid

Signaling Receptor Phospholipase C molecule G protein activated

PIP2

3

Phospholipase C splits PIP2, Activated producing DAG protein kinase C and IP3.

DAG P

P P

GTP

Protein P IP3 P P

P IP3 P P

ER

5 2+

Ca

4

DAG is second messenger that activates protein kinase C enzymes. These enzymes phosphorylate proteins that alter cell processes.

Phosphorylated protein

P

Alters cell activity

IP3 binds to calcium channels in ER. Calcium ions are released into cytosol and act as second messengers.

Alters cell activity

FIGURE 6-10 Phospholipid products as second messengers An activated G protein activates phospholipase C. This enzyme splits PIP2, producing two second messengers—IP3 and DAG. IP3 binds to calcium channels in the endoplasmic reticulum (ER); calcium ions are released into

the cytosol and act as second messengers. DAG activates protein kinase C, a family of enzymes that activates signaling pathways by phosphorylating proteins; calcium ions are needed for the activation of protein kinase C.

rosine in proteins (see Fig. 6-6). When a signaling molecule, such as a growth factor, binds to an enzyme-linked receptor, sites on the receptor are phosphorylated. Addition of the phosphates activates the enzyme part of the receptor. Once activated, tyrosine kinase enzymes can phosphorylate signaling proteins in the cell. Specific target proteins recognize the phosphorylated tyrosines on the tyrosine kinase receptor molecules. These signaling proteins bind to the phosphorylated sites on the receptor molecules. The signaling proteins are then phosphorylated and can activate specific signaling pathways. Recall that in many signal transduction pathways, protein kinases form signaling cascades in which the protein kinases are phosphorylated one after another (see Fig. 6-6). Even when the amino acid phosphorylated in the receptor is tyrosine, the amino acids phosphorylated in the protein kinases in the signaling pathway can be serine or threonine, rather than tyrosine. Each reaction in a chain is dependent on the preceding one. Activation of the last protein in

or regulation of certain ion channels. Protein kinase C stimulates contraction of smooth muscle in the digestive system and in other organs of the body. IP3 is a member of a family of inositol phosphate messengers, some of which can donate phosphate groups to proteins. IP3 binds to calcium channels in the endoplasmic reticulum (ER), causing them to open and release calcium ions into the cytosol. We can summarize this sequence of events as follows: signaling molecule binds to G protein–linked receptor ¡ activates G protein ¡ activates phospholipase ¡ splits PIP2 ¡ inositol trisphosphate (IP3) + diacylglycerol (DAG)

DAG ¡ activates protein kinase enzymes ¡ phosphorylates proteins ¡ some response in cell

IP3 ¡ binds to calcium channels in ER ¡ calcium ions released into the cytosol ¡ some response in the cell

Calcium ions are important messengers Calcium ions (Ca2+) have important functions in many cell processes, including microtubule disassembly, muscle contraction, blood clotting, secretion, and activation of certain cells in the immune system. Calcium ions are critical in neural signaling, including the pathways involved in learning. These ions are also essential in fertilization of an egg and in the initiation of development. Ion pumps in the plasma membrane normally maintain a low calcium ion concentration in the cytosol compared to its concentration in the extracellular fluid. Calcium ions are also stored in the endoplasmic reticulum. When Ca2+ gates open in the plasma membrane or endoplasmic reticulum, the Ca2+ concentration rises in the cytosol. Calcium ions can act alone, but typically they exert their effects by binding to certain proteins. Calmodulin, found in all eukaryotic cells, is an important Ca2+ binding protein. When four Ca2+ bind to a calmodulin molecule, the molecule changes shape and can then activate certain enzymes. Calmodulin combines with a number of different enzymes, including protein kinases and protein phosphatases, and alters their activity. Various types of cells have different calmodulin-binding proteins. How a target cell responds depends on which proteins are present. Calmodulin is important in the regulation of many key processes including metabolism, muscle contraction, memory, inflammation, and apoptosis.

KEY POINT

Many intracellular receptors are transcription factors; when activated, they activate or repress specific genes. Signaling molecules

1

Signaling molecules pass through plasma membrane.

2

3

Signaling molecules move through cytosol.

Signaling molecules pass through nuclear envelope and combine with receptor in nucleus.

Nucleus

Transcription factor (Activated receptor) DNA

4

5

Activated receptor is a transcription factor that activates (or represses) specific genes.

Proteins are synthesized.

Messenger RNA Ribosome Messenger RNA

Protein 6

Cell activity is altered.

Many enzyme-linked receptors activate protein kinase signaling pathways Recall that many enzyme-linked receptors are tyrosine kinases, enzymes that phosphorylate the amino acid ty-

Target cell

FIGURE 6-11 Intracellular receptors

the chain causes a specific cell response. Some cell process is altered. Signaling molecule

Many activated intracellular receptors are transcription factors

Receptor Plasma membrane

Some hydrophobic signaling molecules diffuse across the membranes of target Cytosol cells and bind with intracellular receptors in the cytosol or in the nucleus. For Scaffolding Kinase 1 example, cortisol receptors are located protein in the cytosol. (Cortisol is a steroid horKinase 2 mone produced in the adrenal glands; Kinase 3 its structure is shown in Figure 3-15b.) Thyroid hormones pass into the nucleus and bind with receptors that are bound to DNA in the nucleus. Many intracellular receptors are transcription factors Response that regulate the expression of specific genes. When a signaling molecule binds FIGURE 6-12 A scaffold protein to a receptor, the receptor is activated. Scaffold proteins organize groups of signaling molecules into a signaling complex, making signal transducThe ligand–receptor complex binds to a tion faster, more precise, and more efficient. specific region of DNA and activates or represses specific genes (FIG. 6-11). Cell biologists have demonstrated that when certain signalGene activation can take place quickly—within about 30 mining molecules bind to integrins in the plasma membrane, specific utes. Messenger RNA is produced and carries the code for synthesis signal transduction pathways are activated. Interestingly, growth of a particular protein into the cytoplasm. In combination with rifactors and certain molecules of the extracellular matrix may modbosomes, messenger RNA manufactures specific proteins that can ulate one another’s messages. Integrins also respond to informaalter cell activity. tion received from inside the cell. This inside-out signaling affects how selective integrins are with respect to the molecules to which Scaffold proteins increase efficiency they bind and how strongly they bind to them. Signal transduction is a rapid, precise process. Enzymes must be organized so that they are available as needed for signaling pathReview ways. Scaffold proteins organize groups of intracellular signaling ■ How is an extracellular signal converted to an intracellular signal in molecules into signaling complexes (FIG. 6-12). These proteins signal transduction? Give a specific example. position enzymes close to the proteins they regulate, increasing ■ What is the action of cyclic AMP? of DAG? the probability that they will react with one another. At the same ■ What are the functions of scaffold proteins? time, scaffold proteins decrease the chance that enzymes will be co-opted by other pathways. These organizing proteins guide interactions between molecules, reducing cross talk among different 6.5 RESPONSES TO SIGNALS signaling pathways. Thus, scaffold proteins ensure that signals are relayed accurately, rapidly, and efficiently. ■ ■ LEARNING OBJECTIVES Scaffold proteins have been identified in many pathways, and similar scaffold proteins have been found in diverse organisms. 7 Describe three types of responses that cells make to signaling molecules. Both yeasts and mammals have scaffold proteins that bind kinases 8 Contrast signal amplification with signal termination. in MAP kinase pathways (discussed in the next section).

Signals can be transmitted in more than one direction Integrins, transmembrane proteins that connect the cell to the extracellular matrix, transduce signals in two directions. They transmit signals from outside the cell to the cell interior and also transmit information about the cell interior to the extracellular matrix.

As we have learned, signaling molecules activate signal transduction pathways that bring about specific responses in the cell. Most of these responses fall into three categories: ion channels open or close; enzyme activity is altered, leading to metabolic changes and other effects; and specific gene activity may be turned on or off. Various mechanisms and pathways interact to produce specific actions that are responsible for the structure and function of the cell.

They are responsible for metabolic activity, movement, growth, cell division, development, and further information transfer. In animals, neurons release neurotransmitters that excite or inhibit other neurons or muscle cells by affecting ion channels. For example, when acetylcholine binds with a receptor on a target neuron, an ion channel opens and allows passage of sodium and potassium ions. The resulting change in ion permeability can activate the neuron so that it transmits a neural impulse. Serotonin and some other neurotransmitters work indirectly through G proteins and cyclic AMP. In this chain of events, cAMP activates a kinase that phosphorylates a protein which then closes potassium ion channels. This action leads to transmission of a neural impulse. Some G proteins directly open or close ion channels. Some receptors directly affect enzyme activity, whereas others initiate signal transduction pathways in which enzymes are altered by components of the pathway. When bacteria infect the body, they release certain peptides. Neutrophils, a type of white blood cell, have cell-surface receptors that detect these peptides. When the bacterial peptides bind to a neutrophil’s receptors, enzymes are activated that lead to assembly of microfilaments (actin filaments) and microtubules. Contractions of microfilaments at the far end of the neutrophil force the cytoplasm forward. This action allows the neutrophil to move toward the invading bacteria and destroy them. Microtubules and a variety of proteins, including the contractile protein myosin, appear necessary for this movement. Some signaling molecules affect gene activity. For example, some signaling molecules activate genes that lead to the manufacture of proteins needed for growth and cell division. In both plants and animals, steroid hormones regulate development by causing changes in the expression of specific genes. In animal cells, some steroid hormones bind to nuclear receptors and directly regulate expression of specific target genes. In plant cells, steroid hormones bind to receptors on the cell surface. The signal is then transmitted through a chain of molecules, eventually leading to changes in gene expression. Plant hormones will be discussed in greater detail in Chapter 38, and animal hormones are the focus of Chapter 49.

Ras pathways involve tyrosine kinase receptors and G proteins Some tyrosine kinase receptors activate G proteins. For example, when growth factors bind to tyrosine kinase receptors, ras proteins are activated. Ras proteins are a group of small G proteins that, like other G proteins, are active when bound to GTP. Ras proteins are molecular switches that regulate signaling networks inside the cell. When activated, Ras triggers a cascade of reactions called the Ras pathway. In this pathway, a tyrosine in specific kinase proteins is phosphorylated, leading to critical cell responses. Ras pathways are important in gene expression, cell division, cell movement, cell differentiation, cell adhesion, embryonic development, and apoptosis. For example, to initiate DNA synthesis, fibroblasts (a type of connective tissue cell) require the presence of two growth factors, epidermal growth factor and platelet-derived growth factor. In one study, investigators injected fibroblasts with antibodies that inactivate Ras proteins by binding to them. The

fibroblasts no longer synthesized DNA in response to growth factors. Data from this and similar experiments led to the conclusion that Ras proteins are important in signal transduction involving growth factors. Ras genes code for Ras proteins. Certain mutations in Ras genes result in mutant Ras proteins that bind GTP but cannot hydrolyze it. The mutant Ras proteins are stuck in the “on” state, resulting in unregulated cell division. This condition is associated with several types of human cancer. In fact, mutations in Ras genes have been identified in about one-third of all human cancers. Drugs that inhibit specific tyrosine kinase receptors and Ras pathways are being developed to treat cancer. One Ras pathway that has been extensively studied is the MAP kinase pathway, also known as the ERK pathway; MAP is an acronym for “mitogen-activated protein” (mitogens induce mitosis, the nuclear division associated with eukaryotic cell division). ERK is an acronym for “extracellular signal-regulated kinases.” Several distinct groups of MAP kinases have been described. The pathway illustrated in FIGURE 6-13 shows three main MAP protein kinases: Raf, Mek, and ERK. Proteins in the MAP pathway phosphorylate a nuclear protein that combines with other proteins to form a transcription factor. When specific genes are activated, proteins needed for cell growth, cell division, and cell differentiation (specialization) are synthesized. The MAP kinase cascade is the main signaling pathway for cell division and differentiation. As illustrated in FIGURE 6-14, the signaling proteins ERK 1 and ERK 2 are important in fertility in mammals.

The response to a signal is amplified Signaling molecules are typically present in very low concentrations, yet their effects on the cell are often profound. This is possible because the signal is amplified, as it is relayed through a signaling pathway. For example, let us examine how the action of a signaling molecule such as the hormone epinephrine is magnified as a signal passes through a series of proteins inside the cell. Epinephrine is released by the adrenal glands in response to danger or other stress. Among its many actions, epinephrine increases heart rate, blood flow to skeletal muscle, and glucose concentration in the blood. Epinephrine binds to a G protein–linked receptor, causing the receptor to change shape and activate a G protein. A single molecule of a hormone such as epinephrine can activate many G proteins. Each G protein activates an adenylyl cyclase molecule and then returns to its inactive state. Before it becomes inactive, each adenylyl cyclase can catalyze the production of numerous cAMP molecules (FIG. 6-15). Then, each cAMP molecule can activate many molecules of a particular protein kinase. That protein kinase can phosphorylate many molecules of the next kinase in the pathway and so on down the cascade. As a result of signal amplification, a single signaling molecule can lead to changes in millions of molecules at the end of a signaling cascade. The response is much greater than would be possible if each signaling molecule acted alone. This process of magnifying the strength of a signaling molecule explains how just a few signaling molecules can lead to major responses in the cell.

KEY POINT

When growth factors bind to their receptors, they activate the G protein Ras; Ras activates a MAP-kinase signaling pathway, leading to activation (or repression) of specific genes, and ultimately to protein activity that affects some cell process. 1

Signaling molecule (EGF)

Receptor

Epidermal growth factor (EGF) binds with tyrosine kinase receptor. 2

Inactive Ras

Adapter proteins bind to tyrosine kinase receptor and stimulate removal of GDP from G protein Ras.

3

P

Tyr

Tyr

P

P

Tyr

Tyr

P

P

Tyr

Tyr

P

P

Tyr

Tyr

P

Ras then binds with GTP and becomes active.

Active Ras GDP GTP

GDP Adapter proteins

Raf

ATP

GTP

4

Active Raf

ADP

Active Ras phosphorylates a MAP kinase called Raf, activating it.

P

5

Raf then activates a MAP kinase called Mek.

Mek

ATP

P

ERK 6

Mek activates a MAP kinase called ERK.

7

ERK can activate many different proteins including other protein kinases and transcription factors.

8

Active Mek

ADP

Gene activity is altered, affecting some cell process.

ATP

ADP

Active ERK P

Gene activity altered

Change in some cell process

FIGURE 6-13 A highly simplified Ras/MAP kinase signaling pathway In the pathway illustrated here, epidermal growth factor (EGF) binds with a tyrosine kinase receptor, leading to activation of the small G protein Ras. Then, Ras activates a MAP-kinase signaling pathway. A

Signals must be terminated Once a signal has done its job, it must be terminated. Signal termination returns the receptor and each of the components of the signal transduction pathway to their inactive states. This action allows the magnitude of the response to reflect the strength of the signal. Molecules in the system must also be ready to respond to new signals.

series of MAP kinases in the pathway are activated by phosphorylation. The final MAP kinase can regulate several transcription factors, leading to changes in gene expression that affect cell processes.

We have seen that after a G protein is activated, a subunit of the G protein, a GTPase, catalyzes the hydrolysis of GTP to GDP. This action inactivates the G protein. In the cyclic AMP pathway, any increase in cAMP concentration is temporary. Cyclic AMP is rapidly inactivated by a phosphodiesterase, which converts it to adenosine monophosphate (AMP). Thus, the concentration of cAMP

KEY EXPERIMENT QUESTION: Do the signaling molecules ERK 1 and ERK 2 have key functions in signaling pathways leading to maturation of oocytes (eggs) and ovulation (release of a mature egg from the ovary) in mammals? HYPOTHESIS: The signaling molecules ERK 1 and ERK 2 are key target molecules of the signal sent by the reproductive hormone, luteinizing hormone (LH), and thus are important in oocyte maturation and ovulation in mammals. EXPERIMENT: The researchers produced a line of mice that were deficient in both ERK 1 and ERK 2. They performed biochemical analyses on the signaling pathway activated by LH. They also examined the ovaries of the mice.

LH binds to receptor

depends on the activity of both adenylyl cyclase, which produces it, and of phosphodiesterase, which breaks it down (see Fig. 6-8). Recall also that in many signaling pathways, each protein kinase activates the next protein kinase in the chain by phosphorylating it; a phosphatase then inactivates it by removing the phosphate group. Failure to terminate signals can lead to dire consequences. For example, the bacterium that causes cholera is ingested when people drink contaminated water. Cholera is prevalent in areas where water is contaminated with human feces. The cholera bacterium releases a toxin that activates G proteins in the epithelial cells lining the intestine. The toxin chemically changes the G protein so that it no longer switches off. As a result, the G protein continues to stimulate adenylyl cyclase to make cAMP. The cells lining the intestine malfunction, allowing a large flow of chloride ions into the intestine. Water and other ions follow, leading to the severe watery diarrhea that characterizes cholera. The disease is treated by replacing the lost fluid. If untreated, this G protein malfunction can cause death.

Review

EGF-like factors*

■

What are some cell responses to signals?

■

How do cells amplify signals? What is signal termination?

■

activate

6.6 EVOLUTION OF CELL COMMUNICATION

Ras activates

■ ■ LEARNING OBJECTIVE

ERK 1/ERK 2 activate

9 Cite evidence supporting a long evolutionary history for cell signaling molecules.

Other proteins/factors

Nucleus Activate genes

Oocyte maturation/Ovulation

RESULTS AND CONCUSIONS: In mice deficient in both ERK 1 and ERK 2, the signaling pathway from LH was disrupted. As a result, genes regulating oocyte maturation and ovulation were not activated. Oocytes failed to mature and ovulate, and the mice also exhibited reproductive disorders associated with changes in levels of reproductive hormones. ERK 1 and ERK 2 are key molecules in a signaling pathway that regulates reproduction in mice. These results may lead to greater understanding of certain ovarian disorders that result in infertility in humans. *EGF = epidermal growth factor Source: Fan, H.-Y. et al., Science, Vol. 324, 938–941, 2009.

FIGURE 6-14 Identifying key molecules in a signaling pathway that regulates reproduction in mammals

In this chapter, we have examined how the cells of a multicellular organism signal one another and have described a few of the many signal transduction pathways within cells. We have described quorum sensing and other examples of communication between members of a species. We have also discussed communication among members of different species, such as signaling between plants and insects. In our discussion, we have noted many similarities in the types of signals used and in the molecules that cells use to relay signals from the cell surface to the molecules that carry out a specific response. Some signal transduction pathways found in organisms as diverse as yeasts and animals are quite similar. G proteins, protein kinases, and phosphatases have been highly conserved and are part of signaling pathways in most organisms. Certain disease-causing bacteria have signal transduction pathways similar to those found in eukaryotes. Bacteria may use some of these signal mechanisms to interfere with normal function in the eukaryotic cells they infect. Such similarities in many species suggest evolutionary relationships. Similarities in cell signaling suggest that the molecules and mechanisms used in cell communication are very old. The evidence suggests that cell communication first evolved in prokaryotes and continued to change over time as new types of organisms evolved. However, the fact that some cell signaling molecules have not

Receptor Signaling molecule

1

Adenylyl cyclase activated

One signaling molecule activates many G G protein protein molecules.

GTP 2

GTP

GTP

GTP

Each G protein activates an adenylyl cyclase molecule.

cAMP

ATP cAMP

cAMP cAMP

cAMP

4

Protein kinase Protein 5

Each protein kinase catalyzes phosphorylation of many proteins.

Each adenylyl cyclase catalyzes production of many cAMP molecules.

3

cAMP

Each cAMP molecule activates a protein kinase. P P

P

P

Phosphorylated proteins

P P

P P

6

Phosphorylated proteins affect cell processes.

FIGURE 6-15 Signal amplification The signal is amplified at each step in the pathway so that one activated receptor can give rise to thousands of final products (proteins at the end of the pathway). The response is far greater than what you might expect from a single receptor.

■ ■

changed very much over time suggests that the importance of these pathways to cell survival has restricted any evolutionary changes that might have made them less effective. Thus, these pathways have weathered the demands of natural selection through millions of years of evolution. Choanoflagellates, unicellular protists that are thought to be the most recent ancestors of animals, are used as models for studying the early evolution of animals. Choanoflagellates have many of the same proteins found in animals. These tiny organisms have protein kinases similar to those of animals, and a G protein–linked receptor has been identified. These findings indicate that important proteins necessary for cell communication in animals had evolved long before the evolution of animals. As we will discuss in later chapters, similarities and differences in basic molecules, such as G proteins, can be used to trace evolutionary pathways.

Review ■

Choanoflagellates and animals have similar protein kinases. What does that suggest about their cell signaling mechanisms?

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

6.1 (page 135)

■

1 Describe the four main processes essential for cells to communicate. ■

Cells communicate by cell signaling, which consists of four main processes: (1) synthesis, release, and transport of signaling molecules; (2) reception of information by target cells; (3) signal transduction, the process in which a receptor converts an extracellular signal into an intracellular signal and relays the signal, leading to a cellular response; and (4) response by the cell, for example, some metabolic process may be altered.

6.2 (page 136) 2 Compare three types of signaling molecules: neurotransmitters, hormones, and local regulators Most neurons (nerve cells) signal one another by releasing chemical ■ compounds called neurotransmitters.

■

Hormones are chemical messengers in plants and animals. In animals, they are secreted by endocrine glands, glands that have no ducts. Most hormones diffuse into capillaries and are transported by the blood to target cells. Local regulators diffuse through the interstitial fluid and act on nearby cells. This is called paracrine regulation. Histamine, growth factors, prostaglandins (a type of local hormone), and nitric oxide (a gaseous signaling molecule that passes into target cells) are examples of local regulators.

6.3 (page 137) 3 Identify mechanisms that make reception a highly specific process. ■

Each type of receptor has a specific shape, and only the signaling molecule that fits the specific receptor can affect the cell. A cell can have many different types of receptors and can make different

receptors at different stages in its life cycle or in response to different conditions. Different types of cells can have different types of receptors. 4 Briefly compare ion channel–linked receptors, G protein–linked receptors, enzyme-linked receptors, and intracellular receptors. ■ When a signaling molecule binds to an ion channel–linked receptor, the ion channel opens or, in some cases, closes. G protein–linked receptors are transmembrane proteins that ■ extend into the cytosol or outside the cell. These receptors couple specific signaling molecules to signal transduction pathways inside the cell. The tail of the receptor that extends into the cytosol has a binding site for a specific G protein, a regulatory protein that binds to GTP. ■ Enzyme-linked receptors are transmembrane proteins with a binding site for a signaling molecule outside the cell and a binding site for an enzyme inside the cell. Many enzyme-linked receptors are tyrosine kinases in which the enzyme is part of the receptor.

P

Tyr

Tyr

P

P P P

Tyr

Tyr

Tyr

Tyr

Tyr

Tyr

P P P

Some G proteins directly activate enzymes that catalyze changes in certain proteins, leading to changes in cell function. ■ Intracellular receptors are located in the cytosol or nucleus. These receptors are transcription factors that activate or repress the expression of specific genes. 6 Trace the sequence of events in signal transduction for each of the following second messengers: cyclic AMP, inositol trisphosphate, diacylglycerol, and calcium ions. ■ In many signaling systems, the signaling molecule serves as the first messenger. Information is relayed by the G protein to a second messenger, an intracellular signaling molecule. ■ When certain G proteins undergo a conformational change, they bind with and activate adenylyl cyclase, an enzyme on the cytoplasmic side of the plasma membrane. Adenylyl cyclase catalyzes the formation of cyclic AMP (cAMP) from ATP. Cyclic AMP is a second messenger that activates certain protein kinase enzymes that phosphorylate certain proteins. The phosphorylated protein triggers a chain of reactions that lead to some response in the cell. ■ Certain G proteins activate the membrane-bound enzyme phospholipase C. This enzyme splits a phospholipid, PIP2 (phosphotidylinositol4,5-bisphosphate), into two products, inositol trisphosphate (IP3) and diacylglycerol (DAG). IP3 is a second messenger that can donate phosphate groups to proteins. IP3 binds to calcium channels in the endoplasmic reticulum, which causes the channels to open and release calcium ions into the cytosol. DAG is a second messenger that activates certain protein kinase enzymes. These enzymes phosphorylate a variety of target proteins. ■ Calcium ions can also act as second messengers. They typically combine with the protein calmodulin, which then affects the activity of protein kinases and protein phosphatases.

6.5 (page 147) ■

Intracellular receptors are located in the cytosol or in the nucleus. Their ligands are small, hydrophobic molecules that diffuse across the plasma membrane.

6.4 (page 141) 5 Compare the actions of the main types of receptors in signal transduction. Ion channel–linked receptors convert chemical signals into electrical signals. The gates of many ion channels remain closed until ligands bind to them. ■ Many enzyme-linked receptors are tyrosine kinases, enzymes that phosphorylate proteins. Tyrosine kinase receptors activate several different signal transduction pathways. In a protein kinase cascade, each molecule in the signaling pathway is phosphorylated by the preceding protein kinase in the chain. The last protein kinase in the cascade activates the target protein by phosphorylation. The target protein alters some process in the cell. G protein–linked receptors activate G proteins. A G protein con■ sists of three subunits. It is linked to a molecule of guanosine diphosphate (GDP), a molecule similar to ADP but containing the base guanine instead of adenine. When a ligand binds with the receptor, the GDP is released and is replaced by guanosine triphosphate (GTP). Then one subunit of the G protein separates from the other two subunits. The activated G protein may initiate a signal transduction pathway by binding with a specific protein in the cell. ■

7 Describe three types of responses that cells make to signaling molecules. ■ In response to signaling molecules: ion channels open or close; enzyme activity changes leading to metabolic changes and other effects; and specific genes are activated or repressed. These responses can affect cell shape, cell growth, cell division, cell differentiation, and metabolism. 8 Contrast signal amplification with signal termination. ■ Signal amplification is the process of enhancing the cell’s response to a signal as the signal is relayed through a signal transduction pathway. Before it becomes inactive, each enzyme can catalyze the production of numerous product molecules. ■ Signal termination is the process of inactivating the receptor and each component of the signal transduction pathway once they have done their jobs. Signal termination allows molecules in the system to respond to new signals.

6.6 (page 150) 9 Cite evidence supporting a long evolutionary history for cell signaling molecules. ■ Molecules important in cell signaling first evolved in prokaryotes. G proteins, protein kinases, and phosphatases have been highly conserved and are part of most signaling pathways.

T E S T YO U R U N D E R S TA N D I N G 1. During signal transduction (a) the cell converts an extracellular signal into an intracellular signal that leads to a change in some cell process (b) a signaling molecule activates or represses several genes (c) each enzyme catalyzes production of one molecule of

product (d) relay of the signal is regulated by paracrine regulation (e) the signal is terminated by prostaglandins 2. In paracrine regulation (a) endocrine glands release hormones that are transported by the blood (b) electrical signals are trans-

mitted (c) neurotransmitters signal growth factors (d) prostaglandins terminate signals during signal transduction (e) a local regulator signals nearby cells 3. When a signaling molecule binds with a receptor (a) G proteins are inactivated (b) a third messenger is activated (c) cell signaling is terminated (d) the signaling cell is activated (e) the receptor is activated 4. G protein–linked receptors (a) inactivate G proteins (b) activate first messengers (c) consist of 18 transmembrane alpha helices (d) have a tail that extends into the cytosol and a binding site for a G protein (e) are located in the cytoplasm or nucleus 5. An enzyme-linked receptor (a) is an integral membrane protein (b) would not be found on plant cell surfaces (c) forms a dimer with another enzyme-linked receptor when a ligand binds to it (d) is typically an adenylyl cyclase molecule (e) typically activates ion channels 6. G proteins (a) relay a message from an activated receptor to an enzyme that activates a second messenger (b) are GTP molecules (c) terminate cell signaling (d) directly activate protein kinases (e) function as first messengers

lated (b) the enzyme portion of the receptor is dephosphorylated (c) the receptor is activated and phorphorylates signaling proteins in the cell (d) an ion channel is opened (e) an immediate signal is sent into the nucleus and specific genes are activated or inhibited 11. Scaffold proteins (a) release kinases and phosphatases into the extracellular fluid (b) increase the probability that an enzyme can be used by several different pathways (c) increase accuracy but slow signaling cascades (d) organize groups of intracellular signaling molecules into signaling complexes (e) are transcription factors found mainly in plant cells 12. Each adenylyl cyclase molecule produces many cAMP molecules. This is an example of (a) receptor up-regulation (b) receptor down-regulation (c) signal amplification (d) scaffold (e) similarities produced by evolution 13. Label the following structures in the diagram: first messenger, G protein, ATP, cAMP (second messenger), protein, cell response. (Consult Fig. 6-7 to check your answers.)

7. Which is the correct sequence? 1. protein kinase activated 2. adenylyl cyclase activated 3. cAMP produced 4. proteins phosphorylated 5. G protein activated (a) 1, 2, 3, 5, 4 (b) 5, 3, 2, 1, 4 (c) 5, 2, 3, 4, 1 (d) 5, 2, 3, 1, 4 (e) 2, 3, 1, 4, 5 8. Which is the correct sequence? 1. phospholipase activated 2. G protein activated 3. PIP2 split 4. proteins phosphorylated 5. DAG produced (a) 1, 2, 5, 3, 4 (b) 2, 1, 3, 5, 4 (c) 4, 2, 3, 1, 5 (d) 5, 2, 3, 1, 4 (e) 2, 3, 5, 4, 1

Cytosol

9. Calcium ions (a) can act as second messengers (b) split calmodulin (c) are kept at higher concentration in the cytosol than in the extracellular fluid (d) are produced in the ER by protein kinases and protein phosphatases (e) typically terminate signaling cascades 10. When growth hormone binds to an enzyme-linked receptor (a) G proteins form a cascade of molecules and are phosphory-

CRITICAL THINKING 1. In many instances, the pathway from signaling molecule to final response in the cell involves complex signaling cascades. Give an example of a complex signaling system, and explain why such complexity may have advantages. 2. More than 500 genes have been identified in the human genome that code for protein kinases. What does this imply regarding protein kinases? Explain your answer. 3. EVOLUTION LINK. Cell signaling in plant and animal cells is similar in some ways and different in others. Offer one or more hypotheses for these similarities and differences, and cite specific examples. 4. EVOLUTION LINK. Some of the same G protein–linked receptors and signal transduction pathways found in plants and animals have been identified in fungi and algae. What does this suggest

about the evolution of these molecules? What does it suggest about these molecules and pathways? 5. SCIENCE, TECHNOLOGY, AND SOCIETY. Mutant Ras proteins have been associated with many types of human cancer. Hundreds of millions of dollars are required for the basic research and development of each new drug. As a citizen are you willing to help fund development of new medications to treat cancer through government-sponsored research? Why or why not? If not, what alternatives do you propose? Additional questions are available in CengageNOW at www.cengage.com/ login.

7

Energy and Metabolism

Keren Su/Corbis

Giant panda (Ailuropoda melanoleuca). The chemical energy produced by photosynthesis and stored in bamboo leaves transfers to the panda as it eats.

KEY CONCEPTS

A

7.1 Energy, the capacity to do work, can be kinetic energy

nongrowing cells need energy simply to maintain themselves. Cells ob-

(energy of motion) or potential energy (energy due to position or state).

tain energy in many forms, but that energy can seldom be used directly to

7.2 Energy cannot be created or destroyed (the first law of

energy from one form to another. The ordered systems of the cell provide

thermodynamics), but the total amount of energy available to do work in a closed system decreases over time (the second law of thermodynamics). Organisms do not violate the laws of thermodynamics because, as open systems, they use energy obtained from their surroundings to do work.

7.3 In cells, energy-releasing (exergonic) processes drive energy-requiring (endergonic) processes.

7.4 ATP plays a central role in cell energy metabolism by linking exergonic and endergonic reactions. ATP transfers energy by transferring a phosphate group.

ll living things require energy to carry out life processes. It may seem obvious that cells need energy to grow and reproduce, but even

power cell processes. For this reason, cells have mechanisms that convert the information that makes these energy transformations possible. Because most components of these energy conversion systems evolved very early in the history of life, many aspects of energy metabolism tend to be similar in a wide range of organisms. The sun is the ultimate source of almost all the energy that powers life; this radiant energy flows from the sun as electromagnetic waves. Plants and other photosynthetic organisms capture about 0.02% of the sun’s energy that reaches Earth. As discussed in Chapter 9, photosyn-

7.5 The transfer of electrons in redox reactions is another

thetic organisms convert radiant energy to chemical energy in the bonds

way that cells transfer energy.

of organic molecules. This chemical energy becomes available to plants,

7.6 As biological catalysts, enzymes increase the rate of

animals such as the giant panda shown in the photograph, and other

specific chemical reactions. The activity of an enzyme is influenced by temperature, pH, the presence of cofactors, and inhibitors and/or activators.

organisms through the process of cellular respiration. In cellular respiration, discussed in Chapter 8, organic molecules are broken apart and their energy is converted to more immediately usable forms.

This chapter focuses on some of the basic principles that govern how cells capture, transfer, store, and use energy. We discuss the functions of adenosine triphosphate (ATP) and other molecules used in energy conversions, including those that transfer electrons in oxidation–reduction (redox) reactions. We also pay particular attention to the essential role of enzymes in cell energy dynamics. The flow of energy in ecosystems is discussed in Chapter 55. POTENTIAL Energy of position

7.1 BIOLOGICAL WORK ■ ■ LEARNING OBJECTIVES 1 2

Define energy, emphasizing how it is related to work and to heat. Use examples to contrast potential energy and kinetic energy.

Energy, one of the most important concepts in biology, can be understood in the context of matter, which is anything that has mass and takes up space. Energy is defined as the capacity to do work, which is any change in the state or motion of matter. Technically, mass is a form of energy, which is the basis behind the energy generated by the sun and other stars. More than 4 billion kilograms of matter per second are converted into energy in the sun. Biologists generally express energy in units of work— kilojoules (kJ). It can also be expressed in units of heat energy— kilocalories (kcal)—thermal energy that flows from an object with a higher temperature to an object with a lower temperature. One kilocalorie is equal to 4.184 kJ. Heat energy cannot do cell work because a cell is too small to have regions that differ in temperature. For that reason, the unit most biologists prefer today is the kilojoule. However, we will use both units because references to the kilocalorie are common in the scientific literature.

Organisms carry out conversions between potential energy and kinetic energy When an archer draws a bow, kinetic energy, the energy of motion, is used and work is performed (FIG. 7-1). The resulting tension in the bow and string represents stored, or potential, energy. Potential energy is the capacity to do work as a result of position or state. When the string is released, this potential energy is converted to kinetic energy in the motion of the bow, which propels the arrow. Most actions of an organism involve a series of energy transformations that occur as kinetic energy is converted to potential energy or as potential energy is converted to kinetic energy. Chemical energy, potential energy stored in chemical bonds, is of particular importance to organisms. In our example, the chemical energy of food molecules is converted to kinetic energy in the muscle cells of the archer. The contraction of the archer’s muscles, like many of the activities performed by an organism, is an example of mechanical energy, which performs work by moving matter.

Review ■

You exert tension on a spring and then release it. How do these actions relate to work, potential energy, and kinetic energy?

KINETIC Energy of motion

FIGURE 7-1 Potential versus kinetic energy The potential chemical energy released by cellular respiration is converted to kinetic energy in the muscles, which do the work of drawing the bow. The potential energy stored in the drawn bow is transformed into kinetic energy as the bowstring pushes the arrow toward its target.

7.2 THE LAWS OF THERMODYNAMICS ■ ■ LEARNING OBJECTIVE 3

State the first and second laws of thermodynamics, and discuss the implications of these laws as they relate to organisms.

Thermodynamics, the study of energy and its transformations, governs all activities of the universe, from the life and death of cells to the life and death of stars. When considering thermodynamics, scientists use the term system to refer to an object that they are studying, whether a cell, an organism, or planet Earth. The rest of the universe other than the system being studied constitutes the surroundings. A closed system does not exchange energy with its surroundings, whereas an open system can exchange energy with its surroundings (FIG. 7-2). Biological systems are open systems. Two laws about energy apply to all things in the universe: the first and second laws of thermodynamics.

The total energy in the universe does not change According to the first law of thermodynamics, energy cannot be created or destroyed, although it can be transferred or converted from one form to another, including conversions between matter

Closed system

Open system Energy exchange

Surroundings

Surroundings

(a) A closed system does not exchange energy with its surroundings.

(b) An open system exchanges energy with its surroundings.

FIGURE 7-2 Closed and open systems

and energy. As far as we know, the total mass-energy present in the universe when it formed, almost 14 billion years ago, equals the amount of energy present in the universe today. This is all the energy that can ever be present in the universe. Similarly, the energy of any system plus its surroundings is constant. A system may absorb energy from its surroundings, or it may give up some energy to its surroundings, but the total energy content of that system plus its surroundings is always the same. As specified by the first law of thermodynamics, organisms cannot create the energy they require in order to live. Instead, they must capture energy from the environment and transform it to a form that can be used for biological work.

The entropy of the universe is increasing The second law of thermodynamics states that when energy is converted from one form to another, some usable energy—that is, energy available to do work—is converted into heat that disperses into the surroundings (see Fig. 55-1 for an illustration of energy flow through an ecosystem). As you learned in Chapter 2, heat is the kinetic energy of randomly moving particles. Unlike heat energy, which flows from an object with a higher temperature to one with a lower temperature, this random motion cannot perform work. As a result, the amount of usable energy available to do work in the universe decreases over time. It is important to understand that the second law of thermodynamics is consistent with the first law; that is, the total amount of energy in the universe is not decreasing with time. However, the total amount of energy in the universe that is available to do work is decreasing over time. Less-usable energy is more diffuse, or disorganized. Entropy (S) is a measure of this disorder, or randomness; organized, usable energy has a low entropy, whereas disorganized energy, such as heat, has a high entropy.

Entropy is continuously increasing in the universe in all natural processes. Maybe at some time, billions of years from now, all energy will exist as heat uniformly distributed throughout the universe. If that happens, the universe will cease to operate, because no work will be possible. Everything will be at the same temperature, so there will be no way to convert the thermal energy of the universe into usable mechanical energy. As a consequence of the second law of thermodynamics, no process requiring an energy conversion is ever 100% efficient, because much of the energy is dispersed as heat, increasing entropy. For example, an automobile engine, which converts the chemical energy of gasoline to mechanical energy, is between 20% and 30% efficient. Thus, only 20% to 30% of the original energy stored in the chemical bonds of the gasoline molecules is actually transformed into mechanical energy; the other 70% to 80% dissipates as waste heat. Energy use in your cells is about 40% efficient, with the remaining energy given to the surroundings as heat. Organisms have a high degree of organization, and at first glance they may appear to refute the second law of thermodynamics. As organisms grow and develop, they maintain a high level of order and do not appear to become more disorganized. However, organisms are open systems; they maintain their degree of order over time only with the constant input of energy from their surroundings. This is why plants must photosynthesize and animals must eat. Although the order within organisms may tend to increase temporarily, the total entropy of the universe (organisms plus surroundings) always increases over time.

Review ■ ■

What is the first law of thermodynamics? the second law? Life is sometimes described as a constant struggle against the second law of thermodynamics. How do organisms succeed in this struggle without violating the second law?

7.3 ENERGY AND METABOLISM ■ ■ LEARNING OBJECTIVES 4 Discuss how changes in free energy in a reaction are related to changes in entropy and enthalpy.

5 6

Distinguish between exergonic and endergonic reactions, and give examples of how they may be coupled. Compare the energy dynamics of a reaction at equilibrium with the dynamics of a reaction not at equilibrium.

The chemical reactions that enable an organism to carry on its activities—to grow, move, maintain and repair itself; reproduce; and respond to stimuli—together make up its metabolism. Recall from Chapter 1 that metabolism is the sum of all the chemical activities taking place in an organism. An organism’s metabolism consists of many intersecting series of chemical reactions, or pathways. Two main types of metabolism are anabolism and catabolism. Anabolism includes the various pathways in which complex molecules are synthesized from simpler substances, such as in the linking of amino acids to form proteins. Catabolism includes the pathways in which larger molecules are broken down into smaller ones, such as in the degradation of starch to form monosaccharides.

As you will see, these changes involve not only alterations in the arrangement of atoms but also various energy transformations. Catabolism and anabolism are complementary processes; catabolic pathways involve an overall release of energy, some of which powers anabolic pathways, which have an overall energy requirement. In the following sections, we discuss how to predict whether a particular chemical reaction requires energy or releases it.

Enthalpy is the total potential energy of a system In the course of any chemical reaction, including the metabolic reactions of a cell, chemical bonds break and new and different bonds may form. Every specific type of chemical bond has a certain amount of bond energy, defined as the energy required to break that bond. The total bond energy is essentially equivalent to the total potential energy of the system, a quantity known as enthalpy (H).

Free energy is available to do cell work Entropy and enthalpy are related by a third type of energy, termed free energy (G), which is the amount of energy available to do work under the conditions of a biochemical reaction. (G, also known as “Gibbs free energy,” is named for J. W. Gibbs, a Yale professor who was one of the founders of the science of thermodynamics.) Free energy, the only kind of energy that can do cell work, is the aspect of thermodynamics of greatest interest to a biologist. Enthalpy, free energy, and entropy are related by the following equation: H = G + TS

in which H is enthalpy; G is free energy; T is the absolute temperature of the system, expressed in Kelvin units; and S is entropy. Disregarding temperature for the moment, enthalpy (the total energy of a system) is equal to free energy (the usable energy) plus entropy (the unusable energy). A rearrangement of the equation shows that as entropy increases, the amount of free energy decreases: G = H − TS

Chemical reactions involve changes in free energy Biologists analyze the role of energy in the many biochemical reactions of metabolism. Although the total free energy of a system (G) cannot be effectively measured, the equation G = H − TS can be extended to predict whether a particular chemical reaction will release energy or require an input of energy. The reason

∆G = ∆H − T∆S

Notice that the temperature does not change; it is held constant during the reaction. Thus, the change in free energy (∆G) during the reaction is equal to the change in enthalpy (∆H) minus the product of the absolute temperature (T) in Kelvin units multiplied by the change in entropy (∆S). Scientists express ∆G and ∆H in kilojoules or kilocalories per mole; they express ∆S in kilojoules or kilocalories per Kelvin unit.

Free energy decreases during an exergonic reaction An exergonic reaction releases energy and is said to be a spontaneous or a “downhill” reaction, from higher to lower free energy (FIG. 7-3a). Because the total free energy in its final state is less than the total free energy in its initial state, ∆G is a negative number for exergonic reactions. The term spontaneous may give the false impression that such reactions are always instantaneous. In fact, spontaneous reactions do not necessarily occur readily; some are extremely slow. The reason is that energy, known as activation energy, is required to initiate every reaction, even a spontaneous one. We discuss activation energy later in the chapter.

Free energy increases during an endergonic reaction An endergonic reaction is a reaction in which there is a gain of free energy (FIG. 7-3b). Because the free energy of the products is greater than the free energy of the reactants, ∆G has a positive value. Such a reaction cannot take place in isolation. Instead, it must occur in such a way that energy can be supplied from the

Reactants

Products Free energy decreases

Products

Free energy (G)

Free energy (G)

If we assume that entropy is zero, the free energy is simply equal to the total potential energy (enthalpy); an increase in entropy reduces the amount of free energy. What is the significance of the temperature (T)? Remember that as the temperature increases, there is an increase in random molecular motion, which contributes to disorder and multiplies the effect of the entropy term.

is that changes in free energy can be measured. Scientists use the Greek capital letter delta (∆) to denote any change that occurs in the system between its initial state before the reaction and its final state after the reaction. To express what happens with respect to energy in a chemical reaction, the equation becomes

Free energy increases Reactants

Course of reaction

Course of reaction

(a) In an exergonic reaction, there is a net loss of free energy. The products have less free energy than was present in the reactants, and the reaction proceeds spontaneously.

(b) In an endergonic reaction, there is a net gain of free energy. The products have more free energy than was present in the reactants.

FIGURE 7-3 Animated Exergonic and endergonic reactions

Concentration gradient

A Δ B

Exergonic (process occurs spontaneously)

(a) A concentration gradient is a form of potential energy.

(b) When molecules are evenly distributed, they have high entropy.

FIGURE 7-4 Entropy and diffusion The tendency of entropy to increase can be used to produce work, in this case, diffusion.

surroundings. Of course, many energy-requiring reactions take place in cells, and as you will see, metabolic mechanisms have evolved that supply the energy to “drive” these nonspontaneous cell reactions in a particular direction.

Diffusion is an exergonic process In Chapter 5, you saw that randomly moving particles diffuse down their own concentration gradient (FIG. 7-4). Although the movements of the individual particles are random, net movement of the group of particles seems to be directional. What provides energy for this apparently directed process? A concentration gradient, with a region of higher concentration and another region of lower concentration, is an orderly state. A cell must expend energy to produce a concentration gradient. Because work is done to produce this order, a concentration gradient is a form of potential energy. As the particles move about randomly, the gradient becomes degraded. Thus, free energy decreases as entropy increases. In cellular respiration and photosynthesis, the potential energy stored in a concentration gradient of hydrogen ions (H+) is transformed into chemical energy in adenosine triphosphate (ATP) as the hydrogen ions pass through a membrane down their concentration gradient. This important concept, known as chemiosmosis, is discussed in detail in Chapters 8 and 9.

At the beginning of a reaction, only the reactant molecules (A) may be present. As the reaction proceeds, the concentration of the reactant molecules decreases and the concentration of the product molecules (B) increases. As the concentration of the product molecules increases, they may have enough free energy to initiate the reverse reaction. The reaction thus proceeds in both directions simultaneously; if undisturbed, it eventually reaches a state of dynamic equilibrium, in which the rate of the reverse reaction equals the rate of the forward reaction. At equilibrium there is no net change in the system; a reverse reaction balances every forward reaction. At a given temperature and pressure, each reaction has its own characteristic equilibrium. For any given reaction, chemists can perform experiments and calculations to determine the relative concentrations of reactants and products present at equilibrium. If the reactants have much greater intrinsic free energy than the products, the reaction goes almost to completion; that is, it reaches equilibrium at a point at which most of the reactants have been converted to products. Reactions in which the reactants have much less intrinsic free energy than the products reach equilibrium at a point where very few of the reactant molecules have been converted to products. If you increase the initial concentration of A, then the reaction will “shift to the right,” and more A will be converted to B. A similar effect can be obtained if B is removed from the reaction mixture. The reaction always shifts in the direction that reestablishes equilibrium so that the proportions of reactants and products characteristic of that reaction at equilibrium are restored. The opposite effect occurs if the concentration of B increases or if A is removed; here the system “shifts to the left.” The actual free-energy change that occurs during a reaction is defined mathematically to include these effects, which stem from the relative initial concentrations of reactants and products. Cells use energy to manipulate the relative concentrations of reactants and products of almost every reaction. Cell reactions are virtually never at equilibrium. By displacing their reactions far from equilibrium, cells supply energy to endergonic reactions and direct their metabolism according to their needs.

Free-energy changes depend on the concentrations of reactants and products

Cells drive endergonic reactions by coupling them to exergonic reactions

According to the second law of thermodynamics, any process that increases entropy can do work. As we have discussed, differences in the concentration of a substance, such as between two different parts of a cell, represent a more orderly state than that when the substance is diffused homogeneously throughout the cell. Freeenergy changes in any chemical reaction depend mainly on the difference in bond energies (enthalpy, H) between reactants and products. Free energy also depends on concentrations of both reactants and products. In most biochemical reactions there is little intrinsic freeenergy difference between reactants and products. Such reactions are reversible, indicated by drawing double arrows:

Many metabolic reactions, such as protein synthesis, are anabolic and endergonic. Because an endergonic reaction cannot take place without an input of energy, endergonic reactions are coupled to exergonic reactions. In coupled reactions, the thermodynamically favorable exergonic reaction provides the energy required to drive the thermodynamically unfavorable endergonic reaction. The endergonic reaction proceeds only if it absorbs free energy released by the exergonic reaction to which it is coupled. Consider the free-energy change, ∆G, in the following reaction: (1) A ¡ B

∆G = +20.9 kJ/mol (+5 kcal/mol)

Because ∆G has a positive value, you know that the product of this reaction has more free energy than the reactant. This is an endergonic reaction. It is not spontaneous and does not take place without an energy source. By contrast, consider the following reaction: (2) C ¡ D

∆G = −33.5 kJ/mol (−8 kcal/mol)

The negative value of ∆G tells you that the free energy of the reactant is greater than the free energy of the product. This exergonic reaction proceeds spontaneously. You can sum up reactions 1 and 2 as follows: (1) A ¡ B

∆G = +20.9 kJ/mol (+5 kcal/mol)

(2) C ¡ D

∆G = −33.5 kJ/mol (−8 kcal/mol)

Overall

∆G = −12.6 kJ/mol (−3 kcal/mol)

Because thermodynamics considers the overall changes in these two reactions, which show a net negative value of ∆G, the two reactions taken together are exergonic. The fact that scientists can write reactions this way is a useful bookkeeping device, but it does not mean that an exergonic reaction mysteriously transfers energy to an endergonic “bystander” reaction. However, these reactions are coupled if their pathways are altered so a common intermediate links them. Reactions 1 and 2 might be coupled by an intermediate (I) in the following way: (3) A + C ¡ I

∆G = −8.4 kJ/mol (−2 kcal/mol)

(4) I ¡ B + D

∆G = −4.2 kJ/mol (−1 kcal/mol)

Overall

In all living cells, energy is temporarily packaged within a remarkable chemical compound called adenosine triphosphate (ATP), which holds readily available energy for very short periods. We may think of ATP as the energy currency of the cell. When you work to earn money, you might say your energy is symbolically stored in the money you earn. The energy the cell requires for immediate use is temporarily stored in ATP, which is like cash. When you earn extra money, you may deposit some in the bank; similarly, a cell may deposit energy in the chemical bonds of lipids, starch, or glycogen. Moreover, just as you dare not make less money than you spend, the cell must avoid energy bankruptcy, which would mean its death. Finally, just as you probably do not keep money you earn very long, the cell continuously spends its ATP, which must be replaced immediately. ATP is a nucleotide consisting of three main parts: adenine, a nitrogen-containing organic base; ribose, a five-carbon sugar; and three phosphate groups, identifiable as phosphorus atoms surrounded by oxygen atoms (FIG. 7-5). Notice that the phosphate Adenine NH2 N

N

CH

Oˉ

N

N

H2C O

P

H

H

O

OH

OH

O Ribose

∆G = −12.6 kJ/mol (−3 kcal/mol)

Note that reactions 3 and 4 are sequential. Thus, the reaction pathways have changed, but overall the reactants (A and C) and products (B and D) are the same, and the free-energy change is the same. Generally, for each endergonic reaction occurring in a living cell there is a coupled exergonic reaction to drive it. Often the exergonic reaction involves the breakdown of ATP. Now let’s examine specific examples of the role of ATP in energy coupling.

Phosphate groups

HC

H

H

Oˉ O

Oˉ

˜ P O˜P O

Oˉ

O

Adenosine triphosphate (ATP) Hydrolysis of ATP

H2O

NH2 N

N

CH

Review ■

■

Consider the free-energy change in a reaction in which enthalpy decreases and entropy increases. Is ∆G zero, or does it have a positive value or a negative value? Is the reaction endergonic or exergonic? Why can’t a reaction at equilibrium do work?

HC N

N

Oˉ O H2C

H

H OH

H

O

P O

Oˉ O

˜P

O

■ ■ LEARNING OBJECTIVE 7

Explain how the chemical structure of ATP allows it to transfer a phosphate group and discuss the central role of ATP in the overall energy metabolism of the cell.

OH + HO

P

Oˉ

O

H OH

Adenosine diphosphate (ADP)

7.4 ATP, THE ENERGY CURRENCY OF THE CELL

Oˉ

Inorganic phosphate (Pi )

FIGURE 7-5 Animated ATP and ADP ATP, the energy currency of all living things, consists of adenine, ribose, and three phosphate groups. The hydrolysis of ATP, an exergonic reaction, yields ADP and inorganic phosphate. (The black wavy lines indicate unstable bonds. These bonds allow the phosphates to be transferred to other molecules, making them more reactive.)

groups are bonded to the end of the molecule in a series, rather like three cars behind a locomotive, and, like the cars of a train, they can be attached and detached.

Exergonic reactions release energy

ATP donates energy through the transfer of a phosphate group ADP +

When the terminal phosphate is removed from ATP, the remaining molecule is adenosine diphosphate (ADP) (see Fig. 7-5). If the phosphate group is not transferred to another molecule, it is released as inorganic phosphate (Pi). This is an exergonic reaction with a relatively large negative value of ∆G. (Calculations of the free energy of ATP hydrolysis vary somewhat, but range between about −28 and −37 kJ/mol, or −6.8 to −8.7 kcal/mol.) (5) ATP + H2O ¡ ADP + Pi ∆G = −32 kJ/mol (or −7.6 kcal/mol)

Reaction 5 can be coupled to endergonic reactions in cells. Consider the following endergonic reaction, in which two monosaccharides, glucose and fructose, form the disaccharide sucrose.

(7) glucose + ATP ¡ glucose-P + ADP (8) glucose-P + fructose ¡ sucrose + Pi

Recall from Chapter 6 that a phosphorylation reaction is one in which a phosphate group is transferred to some other compound. In reaction 7 glucose becomes phosphorylated to form glucose phosphate (glucose-P), the intermediate that links the two reactions. Glucose-P, which corresponds to I in reactions 3 and 4, reacts exergonically with fructose to form sucrose. For energy coupling to work in this way, reactions 7 and 8 must occur in sequence. It is convenient to summarize the reactions thus: (9) glucose + fructose + ATP ¡ sucrose + ADP + Pi ∆G = −5 kJ/mol (−1.2 kcal/mol)

When you encounter an equation written in this way, remember that it is actually a summary of a series of reactions and that transitory intermediate products (in this case, glucose-P) are sometimes not shown.

ATP links exergonic and endergonic reactions We have just discussed how the transfer of a phosphate group from ATP to some other compound is coupled to endergonic reactions in the cell. Conversely, adding a phosphate group to adenosine monophosphate, or AMP (forming ADP), or to ADP (forming ATP) requires coupling to exergonic reactions in the cell.

ATP

Energy released drives endergonic reactions

FIGURE 7-6 ATP links exergonic and endergonic reactions Exergonic reactions in catabolic pathways (top) supply energy to drive the endergonic formation of ATP from ADP. Conversely, the exergonic hydrolysis of ATP supplies energy to endergonic reactions in anabolic pathways (bottom).

(6) glucose + fructose ¡ sucrose + H2O ∆G = +27 kJ/mol (or +6.5 kcal/mol)

With a free-energy change of −32 kJ/mol (−7.6 kcal/mol), the hydrolysis of ATP in reaction 5 can drive reaction 6, but only if the reactions are coupled through a common intermediate. The following series of reactions is a simplified version of an alternative pathway that some bacteria use:

Pi

AMP + Pi + energy ¡ ADP ADP + Pi + energy ¡ ATP

Thus, ATP occupies an intermediate position in the metabolism of the cell and is an important link between exergonic reactions, which are generally components of catabolic pathways, and endergonic reactions, which are generally part of anabolic pathways (FIG. 7-6).

The cell maintains a very high ratio of ATP to ADP The cell maintains a ratio of ATP to ADP far from the equilibrium point. ATP constantly forms from ADP and inorganic phosphate as nutrients break down in cellular respiration or as photosynthesis traps the radiant energy of sunlight. At any time, a typical cell contains more than 10 ATP molecules for every ADP molecule. The fact that the cell maintains the ATP concentration at such a high level (relative to the concentration of ADP) makes its hydrolysis reaction even more strongly exergonic and more able to drive the endergonic reactions to which it is coupled. Although the cell maintains a high ratio of ATP to ADP, the cell cannot store large quantities of ATP. The concentration of ATP is always very low, less than 1 mmol/L. In fact, studies suggest that a bacterial cell has no more than a 1-second supply of ATP. Thus, it uses ATP molecules almost as quickly as they are produced. A healthy adult human at rest uses about 45 kg (100 lb) of ATP each day, but the amount present in the body at any given moment is less than 1 g (0.035 oz). Every second in every cell, an estimated 10 million molecules of ATP are made from ADP and phosphate, and an equal number of ATPs transfer their phosphate groups, along with their energy, to whatever chemical reactions need them.

Review ■

Why do coupled reactions typically have common intermediates?

■

■

Give a generalized example of a coupled reaction involving ATP, distinguishing between the exergonic and endergonic reactions. Why is the ATP concentration in a cell about 10 times the concentration of ADP?

7.5 ENERGY TRANSFER IN REDOX REACTIONS ■ ■ LEARNING OBJECTIVE 8

Relate the transfer of electrons (or hydrogen atoms) to the transfer of energy.

You have seen that cells transfer energy through the transfer of a phosphate group from ATP. Energy is also transferred through the transfer of electrons. As discussed in Chapter 2, oxidation is the chemical process in which a substance loses electrons, whereas reduction is the complementary process in which a substance gains electrons. Because electrons released during an oxidation reaction cannot exist in the free state in living cells, every oxidation reaction must be accompanied by a reduction reaction in which the electrons are accepted by another atom, ion, or molecule. Oxidation and reduction reactions are often called redox reactions because they occur simultaneously. The substance that becomes oxidized gives up energy as it releases electrons, and the substance that becomes reduced receives energy as it gains electrons. Redox reactions often occur in a series, as electrons are transferred from one molecule to another. These electron transfers, which are equivalent to energy transfers, are an essential part of cellular respiration, photosynthesis, and many other chemical pro-

cesses. Redox reactions, for example, release the energy stored in food molecules so that ATP can be synthesized using that energy.

Most electron carriers transfer hydrogen atoms Generally, it is not easy to remove one or more electrons from a covalent compound; it is much easier to remove a whole atom. For this reason, redox reactions in cells usually involve the transfer of a hydrogen atom rather than just an electron. A hydrogen atom contains an electron, plus a proton that does not participate in the oxidation–reduction reaction. When an electron, either singly or as part of a hydrogen atom, is removed from an organic compound, it takes with it some of the energy stored in the chemical bond of which it was a part. That electron, along with its energy, is transferred to an acceptor molecule. An electron progressively loses free energy as it is transferred from one acceptor to another. One of the most common acceptor molecules in cellular processes is nicotinamide adenine dinucleotide (NAD+). When NAD+ becomes reduced, it temporarily stores large amounts of free energy. Here is a generalized equation showing the transfer of hydrogen from a compound, which we call X, to NAD+: XH 2 + NAD+¡ X + NADH + H+ Oxidized

Notice that the NAD+ becomes reduced when it combines with hydrogen. NAD+ is an ion with a net charge of +1. When 2 electrons and 1 proton are added, the charge is neutralized and the reduced form of the compound, NADH, is produced (FIG. 7-7). (Although the correct way to write the reduced form of NAD+

NAD+ (oxidized)

NADH (reduced)

H

H

C

HC

CH N

O

Oˉ

P

CH2

O

Phosphate

H

C

HC

CH

C

NH2

+

X

+

H+

O

N

Ribose

H

OH

OH

H

NH2 N

C O

C

HC

CH

C

CH2

H

Adenine

N

N

Phosphate O

O

HC

Nicotinamide

N P

NH2

+

O

H

O

Oˉ

C

C

+

H

H C

HC

H X

Reduced

FIGURE 7-7 NAD+ and NADH O Ribose

H

H

OH

OH

H

NAD+ consists of two nucleotides, one with adenine and one with nicotinamide, that are joined at their phosphate groups. The oxidized form of the nicotinamide ring in NAD+ (left) becomes the reduced form in NADH (right) by the transfer of 2 electrons and 1 proton from another organic compound (XH2), which becomes oxidized (to X) in the process.

is NADH + H+, for simplicity we present the reduced form as NADH in this book.) Some energy stored in the bonds holding the hydrogen atoms to molecule X has been transferred by this redox reaction and is temporarily held by NADH. When NADH transfers the electrons to some other molecule, some of their energy is transferred. This energy is usually then transferred through a series of reactions that ultimately result in the formation of ATP (discussed in Chapter 8). Nicotinamide adenine dinucleotide phosphate (NADP+) is a hydrogen acceptor that is chemically similar to NAD+ but has an extra phosphate group. Unlike NADH, the reduced form of NADP+, abbreviated NADPH, is not involved in ATP synthesis. Instead, the electrons of NADPH are used more directly to provide energy for certain reactions, including certain essential reactions of photosynthesis (discussed in Chapter 9). Other important hydrogen acceptors or electron acceptors are FAD and the cytochromes. Flavin adenine dinucleotide (FAD) is a nucleotide that accepts hydrogen atoms and their electrons; its reduced form is FADH2. The cytochromes are proteins that contain iron; the iron component accepts electrons from hydrogen atoms and then transfers these electrons to some other compound. Like NAD+ and NADP+, FAD and the cytochromes are electron transfer agents. Each exists in a reduced state, in which it has more free energy, or in an oxidized state, in which it has less. Each is an essential component of many redox reaction sequences in cells.

Review ■

Which has the most energy, the oxidized form of a substance or its reduced form? Why?

completed molecule may enter yet another chemical pathway and become totally transformed or consumed to release energy. The changing needs of the cell require a system of flexible metabolic control. The key directors of this control system are enzymes. The catalytic ability of some enzymes is truly impressive. For example, hydrogen peroxide (H2O2) breaks down extremely slowly if the reaction is uncatalyzed, but a single molecule of the enzyme catalase brings about the decomposition of 40 million molecules of hydrogen peroxide per second! Catalase has the highest catalytic rate known for any enzyme. It protects cells by destroying hydrogen peroxide, a poisonous substance produced as a byproduct of some cell reactions. The bombardier beetle uses the enzyme catalase as a defense mechanism (FIG. 7-8).

All reactions have a required energy of activation All reactions, whether exergonic or endergonic, have an energy barrier known as the energy of activation (E A), or activation energy, which is the energy required to break the existing bonds and begin the reaction. In a population of molecules of any kind, some have a relatively high kinetic energy, whereas others have a lower energy content. Only molecules with a relatively high kinetic energy are likely to react to form the product. Even a strongly exergonic reaction, one that releases a substantial quantity of energy as it proceeds, may be prevented from proceeding by the activation energy required to begin the reaction. For example, molecular hydrogen and molecular oxygen can react explosively to form water: 2 H 2 + O 2 ¡ 2 H 2O

7.6 ENZYMES ■ ■ LEARNING OBJECTIVES 9 Explain how an enzyme lowers the required energy of activation for a reaction.

The principles of thermodynamics help us predict whether a reaction can occur, but they tell us nothing about the speed of the reaction. The breakdown of glucose, for example, is an exergonic reaction, yet a glucose solution stays unchanged virtually indefinitely in a bottle if it is kept free of bacteria and molds and not subjected to high temperatures or strong acids or bases. Cells cannot wait for centuries for glucose to break down, nor can they use extreme conditions to cleave glucose molecules. Cells regulate the rates of chemical reactions with enzymes, which are biological catalysts that increase the speed of a chemical reaction without being consumed by the reaction. Although most enzymes are proteins, scientists have learned that some types of RNA molecules have catalytic activity as well (catalytic RNA is discussed in Chapter 13). Cells require a steady release of energy, and they must regulate that release to meet metabolic energy requirements. Metabolic processes generally proceed by a series of steps such that a molecule may go through as many as 20 or 30 chemical transformations before it reaches some final state. Even then, the seemingly

Thomas Eisner and Daniel Aneshansley/Cornell University

10 Describe specific ways enzymes are regulated.

FIGURE 7-8 Catalase as a defense mechanism When threatened, a bombardier beetle (Stenaptinus insignis) uses the enzyme catalase to decompose hydrogen peroxide. The oxygen gas formed in the decomposition ejects water and other chemicals with explosive force. Because the reaction releases a great deal of heat, the water comes out as steam. (A wire attached by a drop of adhesive to the beetle’s back immobilizes it. The researcher prodded its leg with the dissecting needle on the left to trigger the ejection.)

An enzyme lowers a reaction’s activation energy Like all catalysts, enzymes affect the rate of a reaction by lowering the activation energy (EA) necessary to initiate a chemical reaction (FIG. 7-10). If molecules need less energy to react because the activation barrier is lowered, a larger fraction of the reactant molecules reacts at any one time. As a result, the reaction proceeds more quickly. Although an enzyme lowers the activation energy for a reaction, it has no effect on the overall free-energy change; that is, an

KEY POINT

An enzyme lowers the activation energy of a reaction but does not alter the free-energy change.

Activation energy (EA) without enzyme

Free energy (G)

This reaction is spontaneous, yet hydrogen and oxygen can be safely mixed as long as all sparks are kept away because the required activation energy for this particular reaction is relatively high. A tiny spark provides the activation energy that allows a few molecules to react. Their reaction liberates so much heat that the rest react, producing an explosion. Such an explosion occurred on the space shuttle Challenger on January 28, 1986 (FIG. 7-9). The failure of a rubber O-ring to properly seal caused the liquid hydrogen in the tank attached to the shuttle to leak and start burning. When the hydrogen tank ruptured a few seconds later, the resulting force burst the nearby oxygen tank as well, mixing hydrogen and oxygen and igniting a huge explosion.

Activation energy (EA) with enzyme Energy of reactants

Change in free energy (∆G) Energy of products Progress of reaction

FIGURE 7-10 Animated Activation energy and enzymes An enzyme speeds up a reaction by lowering its activation energy (EA). In the presence of an enzyme, reacting molecules require less kinetic energy to complete a reaction.

enzyme can promote only a chemical reaction that could proceed without it. If the reaction goes to equilibrium, no catalyst can cause it to proceed in a thermodynamically unfavorable direction or can influence the final concentrations of reactants and products. Enzymes simply speed up reaction rates.

An enzyme works by forming an enzyme– substrate complex An uncatalyzed reaction depends on random collisions among reactants. Because of its ordered structure, an enzyme reduces this reliance on random events and thereby controls the reaction. The enzyme accomplishes this by forming an unstable intermediate complex with the substrate, the substance on which it acts. When the enzyme–substrate complex, or ES complex, breaks up, the product is released; the original enzyme molecule is regenerated and is free to form a new ES complex: AP/Wide World Photos

enzyme + substrate(s) ¡ ES complex

FIGURE 7-9 The space shuttle Challenger explosion This disaster resulted from an explosive exergonic reaction between hydrogen and oxygen. All seven crew members died in the accident on January 28, 1986.

ES complex ¡ enzyme + product(s)

The enzyme itself is not permanently altered or consumed by the reaction and can be reused. As shown in FIGURE 7-11a, every enzyme contains one or more active sites, regions to which the substrate binds, to form the ES complex. The active sites of some enzymes are grooves or cavities in the enzyme molecule, formed by amino acid side chains. The active sites of most enzymes are located close to the surface. During

Both: Courtesy of Thomas A. Steitz

Active site

(a) Prior to forming an ES complex, the enzyme’s active site is the furrow where the substrate will bind.

(b) The binding of the substrate to the active site induces a change in the conformation of the active site.

FIGURE 7-11 An enzyme–substrate complex This computer graphic model shows the enzyme hexokinase (blue) and its substrate, glucose (red).

the course of a reaction, substrate molecules occupying these sites are brought close together and react with one another. The shape of the enzyme does not seem exactly complementary to that of the substrate. The binding of the substrate to the enzyme molecule causes a change, known as induced fit, in the shape of the enzyme (FIG. 7-11b). Usually, the shape of the substrate also changes slightly, in a way that may distort its chemical bonds. The proximity and orientation of the reactants, together with strains in their chemical bonds, facilitate the breakage of old bonds and the formation of new ones. Thus, the substrate is changed into a product, which diffuses away from the enzyme. The enzyme is then free to catalyze the reaction of more substrate molecules to form more product molecules.

Enzymes are specific Enzymes catalyze virtually every chemical reaction that takes place in an organism. Because the shape of the active site is closely related to the shape of the substrate, most enzymes are highly specific. Most catalyze only a few closely related chemical reactions or, in many cases, only one particular reaction. The enzyme urease, which decomposes urea to ammonia and carbon dioxide, attacks no other substrate. The enzyme sucrase splits only sucrose; it does not act on other disaccharides, such as maltose or lactose. A few enzymes are specific only to the extent that they require the substrate to have a certain kind of chemical bond. For example, lipase, secreted by the pancreas, splits the ester linkages connecting the glycerol and fatty acids of a wide variety of fats. Scientists usually name enzymes by adding the suffix -ase to the name of the substrate. The enzyme sucrase, for example, splits sucrose into glucose and fructose. A few enzymes retain traditional names that do not end in -ase; some of these end in -zyme. For example, lysozyme (from the Greek lysis, “a loosening”) is an enzyme found in tears and saliva; it breaks down bacterial cell walls. Other examples of enzymes with traditional names are pepsin and trypsin, which break peptide bonds in proteins.

Scientists classify enzymes that catalyze similar reactions into groups, although each particular enzyme in the group may catalyze only one specific reaction. TABLE 7-1 describes the six classes of enzymes that biologists recognize. Each class is divided into many subclasses. For example, sucrase, mentioned earlier, is called a glycosidase because it cleaves a glycosidic linkage. Glycosidases are a subclass of the hydrolases (see Fig. 3-8b for the hydrolysis of sucrose). Phosphatases, enzymes that remove phosphate groups by hydrolysis, are also hydrolases. Kinases, enzymes that transfer phosphate groups to substrates, are transferases.

Many enzymes require cofactors Some enzymes consist only of a protein. The enzyme pepsin, which is secreted by the animal stomach and digests dietary protein by breaking certain peptide bonds, is exclusively a protein molecule. Other enzymes have two components: a protein called the apoenzyme and an additional chemical component called a cofactor. Neither the apoenzyme nor the cofactor alone has catalytic activity; only when the two are combined does the enzyme function. A cofactor may be inorganic, or it may be an organic molecule. Some enzymes require a specific metal ion as a cofactor. Two very common inorganic cofactors are magnesium ions and calcium ions. Most of the trace elements, such as iron, copper, zinc, and manganese, all of which organisms require in very small amounts, function as cofactors. An organic, nonpolypeptide compound that binds to the apoenzyme and serves as a cofactor is called a coenzyme. Most coenzymes are carrier molecules that transfer electrons or part of a substrate from one molecule to another. We have already introduced some examples of coenzymes in this chapter. NADH, NADPH, and FADH2 are coenzymes; they transfer electrons.

TABLE 7-1

Important Classes of Enzymes

Enzyme Class

Function

Oxidoreductases Transferases

Catalyze oxidation–reduction reactions Catalyze the transfer of a functional group from a donor molecule to an acceptor molecule Catalyze hydrolysis reactions Catalyze conversion of a molecule from one isomeric form to another Catalyze certain reactions in which two molecules become joined in a process coupled to the hydrolysis of ATP Catalyze certain reactions in which double bonds form or break

Hydrolases Isomerases Ligases

Lyases

ATP functions as a coenzyme; it is responsible for transferring phosphate groups. Yet another coenzyme, coenzyme A, is involved in the transfer of groups derived from organic acids. Most vitamins, which are organic compounds that an organism requires in small amounts but cannot synthesize itself, are coenzymes or components of coenzymes (see descriptions of vitamins in Table 47-3).

Enzymes are most effective at optimal conditions Enzymes generally work best under certain narrowly defined conditions, such as appropriate temperature, pH (FIG. 7-12), and ion concentration. Any departure from optimal conditions adversely affects enzyme activity. Each enzyme has an optimal temperature Most enzymes have an optimal temperature, at which the rate of reaction is fastest. For human enzymes, the temperature optima are near the human body temperature (35°C to 40°C). Enzymatic

Rate of reaction

Most human enzymes

0

Enzymes of heat-tolerant bacteria

10 20 30 40 50 60 70 80 90 100 110 Temperature (°C)

(a) Generalized curves for the effect of temperature on enzyme activity. As temperature increases, enzyme activity increases until it reaches an optimal temperature. Enzyme activity abruptly falls after it exceeds the optimal temperature because the enzyme, being a protein, denatures.

reactions occur slowly or not at all at low temperatures. As the temperature increases, molecular motion increases, resulting in more molecular collisions. The rates of most enzyme-controlled reactions therefore increase as the temperature increases, within limits (see Fig. 7-12a). High temperatures rapidly denature most enzymes. The molecular conformation (3-D shape) of the protein becomes altered as the hydrogen bonds responsible for its secondary, tertiary, and quaternary structures are broken. Because this inactivation is usually not reversible, activity is not regained when the enzyme is cooled. Most organisms are killed by even a short exposure to high temperature; their enzymes are denatured, and they are unable to continue metabolism. There are a few stunning exceptions to this rule. Certain species of archaea (see Chapter 1 for a description of the archaea) can survive in the waters of hot springs, such as those in Yellowstone Park, where the temperature is almost 100°C; these organisms are responsible for the brilliant colors in the terraces of the hot springs (FIG. 7-13). Still other archaea live at temperatures much above that of boiling water, near deep-sea vents, where the extreme pressure keeps water in its liquid state (see Chapter 25 for a discussion of archaea that live in extreme habitats; see also Chapter 55, Inquiring About: Life without the Sun). Each enzyme has an optimal pH Most enzymes are active only over a narrow pH range and have an optimal pH, at which the rate of reaction is fastest. The optimal pH for most human enzymes is between 6 and 8. Recall from Chapter 2 that buffers minimize pH changes in cells so that the pH is maintained within a narrow limit. Pepsin, a protein-digesting enzyme secreted by cells lining the stomach, is an exception; it works only in a very acidic medium, optimally at pH 2 (see Fig. 7-12b). In contrast, trypsin, a protein-splitting enzyme secreted by the pancreas, functions best under the slightly basic conditions found in the small intestine.

Rate of reaction

Trypsin

0

1

2

3

4

5 pH

6

7

8

9

© Henry Holdworth/4wildbynaturegallery.com

Pepsin

10

(b) Enzyme activity is very sensitive to pH. Pepsin is a protein-digesting enzyme in the very acidic stomach juice. Trypsin, secreted by the pancreas into the slightly basic small intestine, digests polypeptides.

FIGURE 7-13 Grand Prismatic Spring in Yellowstone National Park FIGURE 7-12 Animated The effects of temperature and pH on enzyme activity Substrate and enzyme concentrations are held constant in the reactions illustrated.

The world’s third-largest spring, about 61 m (200 ft) in diameter, the Grand Prismatic Spring teems with heat-tolerant archaea. The rings around the perimeter, where the water is slightly cooler, get their distinctive colors from the various kinds of archaea living there.

The activity of an enzyme may be markedly changed by any alteration in pH, which in turn alters electric charges on the enzyme. Changes in charge affect the ionic bonds that contribute to tertiary and quaternary structure, thereby changing the protein’s conformation and activity. Many enzymes become inactive, and usually irreversibly denatured, when the medium is made very acidic or very basic.

Enzymes are organized into teams in metabolic pathways Enzymes play an essential role in reaction coupling because they usually work in sequence, with the product of one enzyme-controlled reaction serving as the substrate for the next. You can picture the inside of a cell as a factory with many different assembly (and disassembly) lines operating simultaneously. An assembly line consists of a number of enzymes. Each enzyme carries out one step, such as changing molecule A into molecule B. Then molecule B is passed along to the next enzyme, which converts it into molecule C, and so on. Such a series of reactions is called a metabolic pathway. Enzyme 1

A

¡ B

Enzyme 2

¡ C

The cell regulates enzymatic activity Enzymes regulate the chemistry of the cell, but what controls the enzymes? One regulatory mechanism involves controlling the amount of enzyme produced. A specific gene directs the synthesis of each type of enzyme. The gene, in turn, may be switched on by a signal from a hormone or by some other signal molecule. When the gene is switched on, the enzyme is synthesized. The total amount of enzyme present then influences the overall cell reaction rate. If the pH and temperature are kept constant (as they are in most cells), the rate of the reaction can be affected by the substrate concentration or by the enzyme concentration. If an excess of substrate is present, the enzyme concentration is the rate-limiting factor. The initial rate of the reaction is then directly proportional to the enzyme concentration (FIG. 7-14a).

Rate of reaction

Rate of reaction

Each of these reactions is reversible, despite the fact that an enzyme catalyzes it. An enzyme does not itself determine the direction of the reaction it catalyzes. However, the overall reaction sequence is portrayed as proceeding from left to right. Recall that if there is little intrinsic free-energy difference between the reactants and products for a particular reaction, the direction of the reaction is determined mainly by the relative concentrations of reactants and products.

In metabolic pathways, both intermediate and final products are often removed and converted to other chemical compounds. Such removal drives the sequence of reactions in a particular direction. Let us assume that reactant A is continually supplied and that its concentration remains constant. Enzyme 1 converts reactant A to product B. The concentration of B is always lower than the concentration of A because B is removed as it is converted to C in the reaction catalyzed by enzyme 2. If C is removed as quickly as it is formed (perhaps by leaving the cell), the entire reaction pathway is “pulled” toward C. In some cases the enzymes of a metabolic pathway bind to one another to form a multienzyme complex that efficiently transfers intermediates in the pathway from one active site to another. An example of one such multienzyme complex, pyruvate dehydrogenase, is discussed in Chapter 8.

Enzyme concentration

(a) In this example, the rate of reaction is measured at different enzyme concentrations, with an excess of substrate present. (Temperature and pH are constant.) The rate of the reaction is directly proportional to the enzyme concentration.

Substrate concentration

(b) In this example, the rate of the reaction is measured at different substrate concentrations, and enzyme concentration, temperature, and pH are constant. If the substrate concentration is relatively low, the reaction rate is directly proportional to substrate concentration. However, higher substrate concentrations do not increase the reaction rate because the enzymes become saturated with substrate.

FIGURE 7-14 The effects of enzyme concentration and substrate concentration on the rate of a reaction

If the enzyme concentration is kept constant, the rate of an enzymatic reaction is proportional to the concentration of substrate present. Substrate concentration is the rate-limiting factor at lower concentrations; the rate of the reaction is therefore directly proportional to the substrate concentration. However, at higher substrate concentrations, the enzyme molecules become saturated with substrate; that is, substrate molecules are bound to all available active sites of enzyme molecules. In this situation, increasing the substrate concentration does not increase the net reaction rate (FIG. 7-14b). The product of one enzymatic reaction may control the activity of another enzyme, especially in a sequence of enzymatic reactions. For example, consider the following metabolic pathway: Enzyme 1 A

Enzyme 2

¡B

¡ C

Enzyme 3

Enzyme 1 (Threonine deaminase) α-Ketobutyrate Enzyme 2 α-Aceto-α-hydroxybutyrate Feedback inhibition

Enzyme 3 α,β-Dihydroxy-β-methylvalerate

(Isoleucine inhibits enzyme 1)

Enzyme 4 α-Keto-β-methylvalerate

Enzyme 4

¡D

Threonine

Enzyme 5

¡E

A different enzyme catalyzes each step, and the final product E may inhibit the activity of enzyme 1. When the concentration of E is low, the sequence of reactions proceeds rapidly. However, an increasing concentration of E serves as a signal for enzyme 1 to slow down and eventually to stop functioning. Inhibition of enzyme 1 stops the entire reaction sequence. This type of enzyme regulation, in which the formation of a product inhibits an earlier reaction in the sequence, is called feedback inhibition (FIG. 7-15). Another method of enzymatic control focuses on the activation of enzyme molecules. In their inactive form, the active sites of the enzyme are inappropriately shaped, so the substrates do not fit. Among the factors that influence the shape of the enzyme are pH, the concentration of certain ions, and the addition of phosphate groups to certain amino acids in the enzyme. Some enzymes have a receptor site, called an allosteric site, on some region of the enzyme molecule other than the active site. (The word allosteric means “another space.”) When a substance binds to an enzyme’s allosteric site, the conformation of the enzyme’s active site changes, thereby modifying the enzyme’s activ-

Isoleucine

FIGURE 7-15 Animated Feedback inhibition Bacteria synthesize the amino acid isoleucine from the amino acid threonine. The isoleucine pathway involves five steps, each catalyzed by a different enzyme. When enough isoleucine accumulates in the cell, the isoleucine inhibits threonine deaminase, the enzyme that catalyzes the first step in this pathway.

ity. Substances that affect enzyme activity by binding to allosteric sites are called allosteric regulators. Some allosteric regulators are allosteric inhibitors that keep the enzyme in its inactive shape. Conversely, the activities of allosteric activators result in an enzyme with a functional active site. The enzyme cyclic AMP-dependent protein kinase is an allosteric enzyme regulated by a protein that binds reversibly to the allosteric site and inactivates the enzyme. Protein kinase is in this inactive form most of the time (FIG. 7-16). When protein kinase activity is

Cyclic AMP Allosteric site Active site

Substrates

Substrates

Regulator (inhibitor)

(a) Inactive form of the enzyme. The enzyme protein kinase is inhibited by a regulatory protein that binds reversibly to its allosteric site. When the enzyme is in this inactive form, the shape of the active site is modified so that the substrate cannot combine with it.

FIGURE 7-16 Animated An allosteric enzyme

(b) Active form of the enzyme. Cyclic AMP removes the allosteric inhibitor and activates the enzyme.

(c) Enzyme–substrate complex. The substrate can then combine with the active site.

Substrate

needed, the compound cyclic AMP (cAMP; see Fig. 3-25 for the structure) contacts the enzyme-inhibitor complex and removes the inhibitory protein, thereby activating the protein kinase. Activation of protein kinases by cAMP is an important aspect of the mechanism of cell signaling, including the action of certain hormones (see Chapters 6 and 49 for discussions of cell signaling).

Enzymes are inhibited by certain chemical agents Most enzymes are inhibited or even destroyed by certain chemical agents. Enzyme inhibition may be reversible or irreversible. Reversible inhibition occurs when an inhibitor forms weak chemical bonds with the enzyme. Reversible inhibition can be competitive or noncompetitive. In competitive inhibition, the inhibitor competes with the normal substrate for binding to the active site of the enzyme (FIG. 7-17a). Usually, a competitive inhibitor is structurally similar to the normal substrate and fits into the active site and combines with the enzyme. However, it is not similar enough to substitute fully for the normal substrate in the chemical reaction, and the enzyme cannot convert it to product molecules. A competitive inhibitor occupies the active site only temporarily and does not permanently damage the enzyme. In competitive inhibition, an active site is occupied by the inhibitor part of the time and by the normal substrate part of the time. If the concentration of the substrate is increased relative to the concentration of the inhibitor, the active site is usually occupied by the substrate. Biochemists demonstrate competitive inhibition experimentally by showing that increasing the substrate concentration reverses competitive inhibition. In noncompetitive inhibition, the inhibitor binds with the enzyme at a site other than the active site (FIG. 7-17b). Such an inhibitor inactivates the enzyme by altering its shape so that the active site cannot bind with the substrate. Many important noncompetitive inhibitors are metabolic substances that regulate enzyme activity by combining reversibly with the enzyme. Allosteric inhibition, discussed previously, is a type of noncompetitive inhibition in which the inhibitor binds to a special site, the allosteric site. In irreversible inhibition, an inhibitor permanently inactivates or destroys an enzyme when the inhibitor combines with one of the enzyme’s functional groups, either at the active site or elsewhere. Many poisons are irreversible enzyme inhibitors. For example, heavy metals such as mercury and lead bind irreversibly to and denature many proteins, including enzymes. Certain nerve gases poison the enzyme acetylcholinesterase, which is important for the functioning of nerves and muscles. Cytochrome oxidase, one of the enzymes that transports electrons in cellular respiration, is especially sensitive to cyanide. Death results from cyanide poisoning because cytochrome oxidase is irreversibly inhibited and no longer transfers electrons from its substrate to oxygen.

Inhibitor Enzyme

Inhibitor binds to active site

Substrate

(a) Competitive inhibition. The inhibitor competes with the normal substrate for the active site of the enzyme. A competitive inhibitor occupies the active site only temporarily.

Substrates

Active site

Inhibitor Enzyme

Active site not suitable for reception of substrates

(b) Noncompetitive inhibition. The inhibitor binds with the enzyme at a site other than the active site, altering the shape of the enzyme and thereby inactivating it.

FIGURE 7-17 Animated Competitive and noncompetitive inhibition

para-aminobenzoic acid (PABA) (FIG. 7-18). When PABA is available, microorganisms can synthesize the vitamin folic acid, which is necessary for growth. Humans do not synthesize folic acid from PABA. For this reason, sulfa drugs selectively affect bacteria. When a sulfa drug is present, the drug competes with PABA for the active site of the bacterial enzyme. When bacteria use the sulfa drug in-

O H2N

C OH Para-aminobenzoic acid (PABA) O

H2N

S

H N

O

R

Generic sulfonamide (Sulfa drug)

Some drugs are enzyme inhibitors

FIGURE 7-18 Para-aminobenzoic acid and sulfonamides

Physicians treat many bacterial infections with drugs that directly or indirectly inhibit bacterial enzyme activity. For example, sulfa drugs have a chemical structure similar to that of the nutrient

Sulfa drugs inhibit an enzyme in bacteria necessary for the synthesis of folic acid, an important vitamin required for growth. (Note the unusual structure of the sulfonamide molecule; sulfur, which commonly forms two covalent bonds, forms six instead.)

stead of PABA, they synthesize a compound that cannot be used to make folic acid. Therefore, the bacterial cells are unable to grow. Penicillin and related antibiotics irreversibly inhibit a bacterial enzyme called transpeptidase. This enzyme establishes some of the chemical linkages in the bacterial cell wall. Bacteria susceptible to these antibiotics cannot produce properly constructed cell walls and are prevented from multiplying effectively. Human cells do not have cell walls and therefore do not use this enzyme. Thus, except for individuals allergic to it, penicillin is harmless to humans. Unfortunately, during the years since it was introduced, resistance to penicillin has evolved in many bacterial strains. The resistant bacteria fight back with an enzyme of their own, penicillinase, which breaks down the penicillin and renders it ineffective. Because bac-

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teria evolve at such a rapid rate, drug resistance is a growing problem in medical practice.

Review What effect does an enzyme have on the required activation energy of a reaction? How does the function of the active site of an enzyme differ from that of an allosteric site? How are temperature and pH optima of an enzyme related to its structure and function? Is allosteric inhibition competitive or noncompetitive?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

7.1 (page 155) 1 Define energy, emphasizing how it is related to work and to heat. Energy is the capacity to do work (expressed in kilojoules, kJ). Energy can be conveniently measured as heat energy, thermal energy that flows from an object with a higher temperature to an object with a lower temperature; the unit of heat energy is the kilocalorie (kcal), which is equal to 4.184 kJ. Heat energy cannot do cell work. 2 Use examples to contrast potential energy and kinetic energy. ■ Potential energy is stored energy; kinetic energy is energy of motion. ■ All forms of energy are interconvertible. For example, photosynthetic organisms capture radiant energy and convert some of it to chemical energy, a form of potential energy that powers many life processes, such as muscle contraction. ■

7.2 (page 155) 3 State the first and second laws of thermodynamics, and discuss the implications of these laws as they relate to organisms. A closed system does not exchange energy with its surroundings. Organisms are open systems that do exchange energy with their surroundings. ■ The first law of thermodynamics states that energy cannot be created or destroyed but can be transferred and changed in form. The first law explains why organisms cannot produce energy; but as open systems, they continuously capture it from the surroundings. ■ The second law of thermodynamics states that disorder (entropy) in the universe, a closed system, is continuously increasing. No energy transfer is 100% efficient; some energy is dissipated as heat, random motion that contributes to entropy (S), or disorder. As open systems, organisms maintain their ordered states at the expense of their surroundings. ■

7.3 (page 156) 4 Discuss how changes in free energy in a reaction are related to changes in entropy and enthalpy. As entropy increases, the amount of free energy decreases, as shown in the equation G = H − TS, in which G is the free energy, H is the enthalpy (total potential energy of the system), T is the absolute temperature (expressed in Kelvin units), and S is entropy. ■ The equation ∆G = ∆H − T∆S indicates that the change in free energy (∆G) during a chemical reaction is equal to the change in enthalpy (∆H) minus the product of the absolute temperature (T) multiplied by the change in entropy (∆S). ■

5 Distinguish between exergonic and endergonic reactions, and give examples of how they may be coupled. ■ An exergonic reaction has a negative value of ∆G; that is, free energy decreases. Such a reaction is spontaneous; it releases free energy that can perform work. ■ Free energy increases in an endergonic reaction. Such a reaction has a positive value of ∆G and is nonspontaneous. In a coupled reaction, the input of free energy required to drive an endergonic reaction is supplied by an exergonic reaction. Learn more about exergonic and endergonic reactions by clicking on the figure in CengageNOW.

6 Compare the energy dynamics of a reaction at equilibrium with the dynamics of a reaction not at equilibrium. ■ When a chemical reaction is in a state of dynamic equilibrium, the rate of change in one direction is exactly the same as the rate of change in the opposite direction; the system can do no work because the free-energy difference between the reactants and products is zero. ■ When the concentration of reactant molecules is increased, the reaction shifts to the right and more product molecules are formed until equilibrium is re-established.

7.4 (page 159) 7 Explain how the chemical structure of ATP allows it to transfer a phosphate group and discuss the central role of ATP in the overall energy metabolism of the cell. Adenosine triphosphate (ATP) is the immediate energy currency ■ of the cell. It donates energy by means of its terminal phosphate group, which is easily transferred to an acceptor molecule. ATP is formed by the phosphorylation of adenosine diphosphate (ADP), an endergonic process that requires an input of energy. ■ ATP is the common link between exergonic and endergonic reactions and between catabolism (degradation of large complex molecules into smaller, simpler molecules) and anabolism (synthesis of complex molecules from simpler molecules).

7.5 (page 161) 8 Relate the transfer of electrons (or hydrogen atoms) to the transfer of energy. Energy is transferred in oxidation–reduction (redox) reactions. A ■ substance becomes oxidized as it gives up one or more electrons to another substance, which becomes reduced in the process. Electrons are commonly transferred as part of hydrogen atoms.

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NAD+ and NADP+ accept electrons as part of hydrogen atoms and become reduced to form NADH and NADPH, respectively. These electrons (along with some of their energy) can be transferred to other acceptors.

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7.6 (page 162) 9 Explain how an enzyme lowers the required energy of activation for a reaction. An enzyme is a biological catalyst; it greatly increases the speed of a chemical reaction without being consumed.

An enzyme works by lowering the activation energy (EA), the energy necessary to get a reaction going. The active site of an enzyme is a 3-D region where substrates come into close contact and thereby react more readily. When a substrate binds to an active site, an enzyme–substrate complex forms in which the shapes of the enzyme and substrate change slightly. This induced fit facilitates the breaking of bonds and formation of new ones. Learn more about activation energy by clicking on the figure in CengageNOW.

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10 Describe specific ways enzymes are regulated. ■ ■

Activation energy (EA) without enzyme

Free energy (G)

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Activation energy (EA) with enzyme Energy of reactants

Change in free energy (∆G) Energy of products Progress of reaction

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Enzymes work best at specific temperature and pH conditions. A cell can regulate enzymatic activity by controlling the amount of enzyme produced and by regulating metabolic conditions that influence the shape of the enzyme. Some enzymes have allosteric sites, noncatalytic sites to which an allosteric regulator binds, changing the enzyme’s activity. Some allosteric enzymes are subject to feedback inhibition, in which the formation of an end product inhibits an earlier reaction in the metabolic pathway. Reversible inhibition occurs when an inhibitor forms weak chemical bonds with the enzyme. Reversible inhibition may be competitive, in which the inhibitor competes with the substrate for the active site, or noncompetitive, in which the inhibitor binds with the enzyme at a site other than the active site. Irreversible inhibition occurs when an inhibitor combines with an enzyme and permanently inactivates it. Learn more about enzymes by clicking on the figures in CengageNOW.

T E S T YO U R U N D E R S TA N D I N G 1. Which of the following can do work in a cell? (a) entropy (b) heat (c) heat energy (d) all of the above (e) none of the above 2. In a chemical reaction occurring in a cell, free energy is equivalent to (a) heat energy (b) heat (c) disorder (d) potential energy (e) more than one of the above is true 3. Cells are able to function because they (a) are subject to the laws of thermodynamics (b) have mechanisms that transform energy from the environment into useful forms (c) can use enzymes to convert endergonic reactions into spontaneous reactions (d) all of the above 4. Diffusion is an (a) endergonic process because free energy increases (b) endergonic process because free energy decreases (c) exergonic process because entropy increases (d) exergonic process because entropy decreases (e) more than one of the above 5. A spontaneous reaction is one in which the change in free energy (∆G) has a _____ value. (a) positive (b) negative (c) positive or negative (d) none of these (∆G has no measurable value)

(a) A ¡ B, ∆G = +6.08 kJ/mol (b) C ¡ D, ∆G = +3.56 kJ/mol (c) E ¡ F, ∆G = 0 kJ/mol (d) G ¡ H, ∆G = −1.22 kJ/mol (e) I ¡ J, ∆G = −5.91 kJ/mol 8. Consider this reaction: glucose + 6 O2 ¡ 6 CO2 + 6 H2O (∆G = −2880 kJ/mol). Which of the following statements about this reaction is not true? (a) the reaction is spontaneous in a thermodynamic sense (b) a small amount of energy (activation energy) must be supplied to start the reaction, which then proceeds with a release of energy (c) the reaction is exergonic (d) the reaction can be coupled to an endergonic reaction (e) the reaction must be coupled to an exergonic reaction 9. The required energy of activation of a reaction (a) is fixed, and cannot be altered (b) can be lowered by a specific enzyme (c) can be raised by a specific enzyme (d) b or c, depending on the enzyme (e) none of the above

6. Healthy living cells maintain (a) ATP and ADP at equilibrium (b) equal concentrations of ATP and ADP (c) an ATP/ADP ratio of at least 10:1 (d) an ATP/ADP ratio of no more than 1:10 (e) most of the cell’s stored energy in the form of ATP

10. “Induced fit” means that when a substrate binds to an enzyme’s active site (a) it fits perfectly, like a key in a lock (b) the substrate and enzyme undergo conformational changes (c) a site other than the active site undergoes a conformational change (d) the substrate and the enzyme become irreversibly bound to each other (e) c and d

7. Which of the following reactions could be coupled to an endergonic reaction with ∆G = +3.56 kJ/mol?

11. The function of a biochemical pathway is to (a) supply energy to reactions (b) drive a sequence of reactions in a particular direc-

tion (c) maintain chemical equilibrium (d) make energy available to endergonic reactions (e) any of the above, depending on the pathway 12. In the following reaction series, which enzyme(s) is/are most likely to have an allosteric site to which the end product E binds?

(a) enzyme 1 (b) enzyme 2 (c) enzyme 3 (d) enzyme 4 (e) enzymes 3 and 4 Enzyme 1 A

¡B

Enzyme 2

¡ C

Enzyme 3

¡D

Enzyme 4

¡E

CRITICAL THINKING 1. Given what you have learned in this chapter, explain why an extremely high fever (body temperature above 40°C, or 105°F) can be fatal. 2. EVOLUTION LINK. What does the fact that all organisms use ATP/ADP as central links between exergonic and endergonic reactions suggest about the evolution of energy metabolism? 3. EVOLUTION LINK. Some have argued that “evolution is impossible because the second law of thermodynamics states that entropy always increases; therefore natural processes cannot give rise to greater complexity.” In what ways is this statement a misunderstanding of the laws of thermodynamics? 4. ANALYZING DATA. Does the figure illustrate an exergonic reaction or an endergonic reaction? How do you know?

Free energy (G)

Reactants Free energy decreases

Products Course of reaction

5. ANALYZING DATA. Reactions 1 and 2 happen to have the same standard free-energy change: ∆G − 41.8 kJ/mol (−10 kcal/ mol). Reaction 1 is at equilibrium, but reaction 2 is far from equilibrium. Is either reaction capable of performing work? If so, which one? 6. ANALYZING DATA. You are performing an experiment in which you are measuring the rate at which succinate is converted to fumarate by the enzyme succinic dehydrogenase. You decide to add a little malonate to make things interesting. You observe that the reaction rate slows markedly and hypothesize that malonate is inhibiting the reaction. Design an experiment that will help you decide whether malonate is acting as a competitive inhibitor or a noncompetitive inhibitor. Additional questions are available in CengageNOW at www.cengage.com/ login.

8

How Cells Make ATP: Energy-Releasing Pathways

Digital Vision/Getty Images

Grizzly bear (Ursus arctos). This grizzly, shown attempting to eat a jumping salmon, may also eat fruit, nuts, roots, insects, and small vertebrates such as mice and ground squirrels.

KEY CONCEPTS

C

ells are tiny factories that process materials on the molecular level, through thousands of metabolic reactions. Cells exist in a dynamic

8.1 Aerobic respiration is an exergonic redox process in which glucose becomes oxidized, oxygen becomes reduced, and energy is captured to make ATP.

state and are continuously building up and breaking down the many

8.2 Aerobic respiration consists of four stages: glycolysis,

splitting complex molecules into smaller components, and anabolism,

formation of acetyl coenzyme A, the citric acid cycle, and the electron transport chain and chemiosmosis.

8.3 Nutrients other than glucose, including many carbohy-

different cell constituents. As you learned in Chapter 7, metabolism has two complementary components: catabolism, which releases energy by the synthesis of complex molecules from simpler building blocks. Anabolic reactions produce proteins, nucleic acids, lipids, polysaccharides,

drates, lipids, and amino acids, can be oxidized by aerobic respiration.

and other molecules that help maintain the cell or the organism. Most

8.4 Anaerobic respiration and fermentation are ATPyielding redox processes in which glucose becomes oxidized, but oxygen does not become reduced. Instead, these processes involve the reduction of inorganic substances (in anaerobic respiration) or organic substances (in fermentation).

source to drive them.

anabolic reactions are endergonic and require ATP or some other energy Every organism must extract energy from food molecules that it either manufactures by photosynthesis or obtains from the environment. Grizzly bears, such as the one in the photograph, obtain organic molecules from their varied plant and animal diets. How do they obtain energy from these organic molecules? First, the food molecules are broken down by digestion into simpler components that are absorbed into the blood and transported to all the cells. The catabolic processes that convert the energy in the chemical bonds of nutrients to chemical energy stored in ATP then occur inside cells, usually through a process known as cellular respiration.

Cellular respiration may be either aerobic or anaerobic. Aerobic

Oxidation

C6H12 O6 ⫹ 6 O2

respiration requires oxygen, whereas anaerobic pathways, which

6 CO2 ⫹ 6 H2O ⫹ energy (in the chemical bonds of ATP)

include anaerobic respiration and fermentation, do not require oxy-

Reduction

gen. In the process of organismal respiration (discussed in Chapter

If we analyze this summary reaction, it appears that CO2 is produced by the removal of hydrogen atoms from glucose. Conversely, water seems to be formed as oxygen accepts the hydrogen atoms. Because the transfer of hydrogen atoms is equivalent to the transfer of electrons, this is a redox reaction in which glucose becomes oxidized and oxygen becomes reduced (see discussion of redox reactions in Chapters 2 and 7). The products of the reaction would be the same if the glucose were simply placed in a test tube and burned in the presence of oxygen. However, if a cell were to burn glucose, its energy would be released all at once as heat, which not only would be unavailable to the cell but also would actually destroy it. For this reason, cells do not transfer hydrogen atoms directly from glucose to oxygen. Aerobic respiration includes a series of redox reactions in which electrons associated with the hydrogen atoms in glucose are transferred to oxygen in a series of steps (FIG. 8-1). During this process, the free energy of the electrons is coupled to ATP synthesis.

46), your lungs provide a steady supply of oxygen that enables your cells to capture energy through aerobic respiration, which is by far the most common pathway and the main subject of this chapter. All three pathways—aerobic respiration, anaerobic respiration, and fermentation—are exergonic and release free energy that can be captured by the cell.

8.1 REDOX REACTIONS ■ ■ LEARNING OBJECTIVE 1

Write a summary reaction for aerobic respiration that shows which reactant becomes oxidized and which becomes reduced.

Most eukaryotes and prokaryotes carry out aerobic respiration, a form of cellular respiration requiring molecular oxygen (O2). During aerobic respiration, nutrients are catabolized to carbon dioxide and water. Most cells use aerobic respiration to obtain energy from glucose, which enters the cell through a specific transport protein in the plasma membrane (see discussion of facilitated diffusion in Chapter 5). The overall reaction pathway for the aerobic respiration of glucose is summarized as follows:

Review ■

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In the overall reaction of aerobic respiration, which reactant becomes oxidized and which becomes reduced? What is the specific role of oxygen in most cells?

8.2 THE FOUR STAGES OF AEROBIC RESPIRATION

C6H12O6 + 6 O2 + 6 H2O ¡ 6 CO2 + 12 H2O + energy (in the chemical bonds of ATP)

■ ■ LEARNING OBJECTIVES

Note that water is shown on both sides of the equation because it is a reactant in some reactions and a product in others. For purposes of discussion, the equation for aerobic respiration can be simplified to indicate that there is a net yield of water:

2 3

List and give a brief overview of the four stages of aerobic respiration. Indicate where each stage of aerobic respiration takes place in a eukaryotic cell.

e1 e2

E

e3

E = e1 + e2 + e3 + e4 + e5 e4 e5

FIGURE 8-1 Changes in free energy The release of energy from a glucose molecule is analogous to the liberation of energy by a falling object. The total energy released (E) is the same whether it occurs all at once or in a series of steps.

also carry out these processes, but because prokaryotic cells lack mitochondria, the reactions of aerobic respiration occur in the cytosol and in association with the plasma membrane.

4 Add up the energy captured (as ATP, NADH, and FADH2) in each stage of 5 6

aerobic respiration. Define chemiosmosis and explain how a gradient of protons is established across the inner mitochondrial membrane. Describe the process by which the proton gradient drives ATP synthesis in chemiosmosis.

1. Glycolysis. A six-carbon glucose molecule is converted to two three-carbon molecules of pyruvate.1 Some of the energy of glucose is captured with the formation of two kinds of energy

The chemical reactions of the aerobic respiration of glucose are grouped into four stages (FIG. 8-2 and TABLE 8-1; see also the summary equations at the end of the chapter). In eukaryotes, the first stage (glycolysis) takes place in the cytosol, and the remaining stages take place inside mitochondria. Most bacteria and archaea

KEY POINT

1 Pyruvate and many other compounds in cellular respiration exist as anions at the pH found in the cell. They sometimes associate with H+ to form acids. For example, pyruvate forms pyruvic acid. In some textbooks these compounds are presented in the acid form.

The stages of aerobic respiration occur in specific locations.

1

2

3

4

Glycolysis

Formation of acetyl coenzyme A

Citric acid cycle

Electron transport and chemiosmosis

Glucose Mitochondrion Acetyl coenzyme A

Citric acid cycle

Electron transport and chemiosmosis

Pyruvate

2 ATP

2 ATP

32 ATP

FIGURE 8-2 Animated The four stages of aerobic respiration Glycolysis, the first stage of aerobic respiration, occurs in the cytosol. Pyruvate, the product of glycolysis, enters a mitochondrion, where cellular respiration continues with the formation of acetyl CoA, the citric acid cycle, and electron transport and chemiosmosis. Most ATP is synthesized by chemiosmosis.

TABLE 8-1

Summary of Aerobic Respiration

Stage

Summary

Some Starting Materials

Some End Products

1. Glycolysis (in cytosol)

Series of reactions in which glucose is degraded to pyruvate; net profit of 2 ATPs; hydrogen atoms are transferred to carriers; can proceed anaerobically

Glucose, ATP, NAD+, ADP, Pi

Pyruvate, ATP, NADH

2. Formation of acetyl CoA (in mitochondria)

Pyruvate is degraded and combined with coenzyme A to form acetyl CoA; hydrogen atoms are transferred to carriers; CO2 is released

Pyruvate, coenzyme A, NAD+

Acetyl CoA, CO2, NADH

3. Citric acid cycle (in mitochondria)

Series of reactions in which the acetyl portion of acetyl CoA is degraded to CO2; hydrogen atoms are transferred to carriers; ATP is synthesized

Acetyl CoA, H2O, NAD+, FAD, ADP, Pi

CO2, NADH, FADH2, ATP

4. Electron transport and chemiosmosis (in mitochondria)

Chain of several electron transport molecules; electrons are passed along chain; released energy is used to form a proton gradient; ATP is synthesized as protons diffuse down the gradient; oxygen is final electron acceptor

NADH, FADH2, O2, ADP, Pi

ATP, H2O, NAD+, FAD

carriers, ATP and NADH.2 See Chapter 7 to review how ATP transfers energy by transferring a phosphate group (see Figs. 7-5 and 7-6). NADH is a reduced molecule that transfers energy by transferring electrons as part of a hydrogen atom (see Fig. 7-7). 2. Formation of acetyl coenzyme A. Each pyruvate enters a mitochondrion and is oxidized to a two-carbon group (acetate) that combines with coenzyme A, forming acetyl coenzyme A. NADH is produced, and carbon dioxide is released as a waste product. 3. The citric acid cycle. The acetate group of acetyl coenzyme A combines with a four-carbon molecule (oxaloacetate) to form a six-carbon molecule (citrate). In the course of the cycle, citrate is recycled to oxaloacetate, and carbon dioxide is released as a waste product. Energy is captured as ATP and the reduced, high-energy compounds NADH and FADH2 (see Chapter 7 to review FADH2). 4. Electron transport and chemiosmosis. The electrons removed from glucose during the preceding stages are transferred from NADH and FADH2 to a chain of electron acceptor compounds. As the electrons are passed from one electron acceptor to another, some of their energy is used to transport hydrogen ions (protons) across the inner mitochondrial membrane, forming a proton gradient. In a process known as chemiosmosis (described later), the energy of this proton gradient is used to produce ATP. Most reactions involved in aerobic respiration are one of three types: dehydrogenations, decarboxylations, and those we informally categorize as preparation reactions. Dehydrogenations are reactions in which two hydrogen atoms (actually, 2 electrons plus 1 or 2 protons) are removed from the substrate and transferred to NAD+ or FAD. Decarboxylations are reactions in which part of a carboxyl group (¬COOH) is removed from the substrate as a molecule of CO2. The carbon dioxide you exhale with each breath is derived from decarboxylations that occur in your cells. The rest of the reactions are preparation reactions in which molecules undergo rearrangements and other changes so that they can undergo further dehydrogenations or decarboxylations. As you examine the individual reactions of aerobic respiration, you will encounter these three basic types. In following the reactions of aerobic respiration, it helps to do some bookkeeping as you go along. Because glucose is the starting material, it is useful to express changes on a per glucose basis. We will pay particular attention to changes in the number of carbon atoms per molecule and to steps in which some type of energy transfer takes place.

In glycolysis, glucose yields two pyruvates The word glycolysis comes from Greek words meaning “sugar splitting,” which refers to the fact that the sugar glucose is metabolized. Glycolysis does not require oxygen and proceeds under aerobic or anaerobic conditions. FIGURE 8-3 shows a simplified overview of glycolysis, in which a glucose molecule consisting of six carbons is Although the correct way to write the reduced form of NAD+ is NADH + H+, for simplicity we present the reduced form as NADH throughout the book.

2

converted to two molecules of pyruvate, a three-carbon molecule. Some of the energy in the glucose is captured; there is a net yield of two ATP molecules and two NADH molecules. The reactions of glycolysis take place in the cytosol, where the necessary reactants, such as ADP, NAD+, and inorganic phosphate, float freely and are used as needed. The glycolysis pathway consists of a series of reactions, each of which is catalyzed by a specific enzyme (FIG. 8-4, pages 178 and 179). Glycolysis is divided into two major phases: the first includes endergonic reactions that require ATP, and the second includes exergonic reactions that yield ATP and NADH. The first phase of glycolysis requires an investment of ATP The first phase of glycolysis is sometimes called the “energy investment phase” (see Fig. 8-4, steps l to 5). Glucose is a relatively stable molecule and is not easily broken down. In two separate phosphorylation reactions, a phosphate group is transferred from ATP to the sugar. The resulting phosphorylated sugar (fructose1,6-bisphosphate) is less stable and is broken enzymatically into two three-carbon molecules, dihydroxyacetone phosphate and glyceraldehyde-3-phosphate (G3P). The dihydroxyacetone phosphate is enzymatically converted to G3P, so the products at this point in glycolysis are two molecules of G3P per glucose. We can summarize this portion of glycolysis as follows: glucose + 2 ATP Six-carbon compound

2 G3P + 2 ADP Threecarbon compound

The second phase of glycolysis yields NADH and ATP The second phase of glycolysis is sometimes called the “energy capture phase” (see Fig. 8-4, steps 6 to 10). Each G3P is converted to pyruvate. In the first step of this process, each G3P is oxidized by the removal of 2 electrons (as part of two hydrogen atoms). These immediately combine with the hydrogen carrier molecule, NAD+: NAD+ + Oxidized

2 H ¡ NADH + H+

(From G3P)

Reduced

Because there are two G3P molecules for every glucose, two NADH are formed. The energy of the electrons carried by NADH is used to form ATP later. This process is discussed in conjunction with the electron transport chain. In two of the reactions leading to the formation of pyruvate, ATP forms when a phosphate group is transferred to ADP from a phosphorylated intermediate (see Fig. 8-4, steps 7 and l0). This process is called substrate-level phosphorylation. Note that in the energy investment phase of glycolysis two molecules of ATP are consumed, but in the energy capture phase four molecules of ATP are produced. Thus, glycolysis yields a net energy profit of two ATPs per glucose. We can summarize the energy capture phase of glycolysis as follows: 2 G3P + 2 NAD+ + 4 ADP ¡ 2 pyruvate + 2 NADH + 4 ATP

KEY POINT

Glycolysis includes both energy investment and energy capture.

Glycolysis

Formation of acetyl coenzyme A

Citric acid cycle

Electron transport and chemiosmosis

Glucose

GLYCOLYSIS Energy investment phase and splitting of glucose Two ATPs invested per glucose

Pyruvate

2 ATP

2 ATP

32 ATP

Glucose

2 ATP 3 steps 2 ADP

Fructose-1,6-bisphosphate P

P

Glyceraldehyde phosphate (G3P)

Glyceraldehyde phosphate (G3P)

P

P

Energy capture phase Four ATPs and two NADH produced per glucose P (G3P)

P (G3P)

NAD+

NAD+

NADH

NADH 5 steps

2 ADP

2 ADP

2 ATP

2 ATP

Pyruvate

Pyruvate

Net yield per glucose: Two ATPs and two NADH

FIGURE 8-3 Animated An overview of glycolysis The black spheres represent carbon atoms. The energy investment phase of glycolysis leads to the splitting of sugar; ATP and NADH are produced during the energy capture phase. During glycolysis, each glucose molecule is converted to two pyruvates, with a net yield of two ATP molecules and two NADH molecules.

Pyruvate is converted to acetyl CoA In eukaryotes, the pyruvate molecules formed in glycolysis enter the mitochondria, where they are converted to acetyl coenzyme A (acetyl CoA). These reactions occur in the cytosol of aerobic prokaryotes. In this series of reactions, pyruvate undergoes a process known as oxidative decarboxylation. First, a carboxyl group is removed as carbon dioxide, which diffuses out of the cell (FIG. 8-5). Then the remaining two-carbon fragment becomes oxidized, and NAD+ accepts the electrons removed during the oxidation. Finally, the oxidized two-carbon fragment, an acetyl group, becomes attached to coenzyme A, yielding acetyl CoA. Pyruvate dehydrogenase, the enzyme that catalyzes these reactions, is an enormous multienzyme complex consisting of 72 polypeptide chains! Recall from Chapter 7 that coenzyme A transfers groups derived from organic acids. In this case, coenzyme A transfers an acetyl group, which is related to acetic acid. Coenzyme A is manufactured in the cell from one of the B vitamins, pantothenic acid. The overall reaction for the formation of acetyl coenzyme A is: 2 pyruvate + 2 NAD+ + 2 CoA ¡ 2 acetyl CoA + 2 NADH + 2 CO2

Note that the original glucose molecule has now been partially oxidized, yielding two acetyl groups and two CO2 molecules. The electrons removed have reduced NAD+ to NADH. At this point in aerobic respiration, four NADH molecules have been formed as a result of the catabolism of a single glucose molecule: two during glycolysis and two during the formation of acetyl CoA from pyruvate. Keep in mind that these NADH molecules will be used later (during electron transport) to form additional ATP molecules.

The citric acid cycle oxidizes acetyl CoA The citric acid cycle is also known as the tricarboxylic acid (TCA) cycle and the Krebs cycle, after Hans Krebs, the German biochemist who assembled the accumulated contributions of many scientists and worked out the details of the cycle in the 1930s. He received a Nobel Prize in Physiology or Medicine in 1953 for this contribution. A simplified overview of the citric acid cycle, which takes place in the matrix of the mitochondria, is given in FIGURE 8-6, page 180. The eight steps of the citric acid cycle are shown in FIGURE 8-7, page 181. A specific enzyme catalyzes each reaction. The first reaction of the cycle occurs when acetyl CoA transfers its two-carbon acetyl group to the four-carbon acceptor compound oxaloacetate, forming citrate, a six-carbon compound. oxaloacetate + acetyl CoA

citrate + CoA

Four -carbon compound

Six-carbon compound

Two-carbon compound

The citrate then goes through a series of chemical transformations, losing first one and then a second carboxyl group as CO2. One ATP is formed (per acetyl group) by substrate-level phosphorylation. Most of the energy made available by the oxidative steps of the cycle is transferred as energy-rich electrons to NAD+, forming NADH. For each acetyl group that enters the citric acid cycle, three molecules of NADH are produced (steps 3, 4, and 8). Electrons are also transferred to the electron acceptor FAD, forming FADH2.

In the course of the citric acid cycle, two molecules of CO2 and the equivalent of eight hydrogen atoms (8 protons and 8 electrons) are removed, forming three NADH and one FADH2. You may wonder why more hydrogen equivalents are generated by these reactions than entered the cycle with the acetyl CoA molecule. These hydrogen atoms come from water molecules that are added during the reactions of the cycle. The CO2 produced accounts for the two carbon atoms of the acetyl group that entered the citric acid cycle. At the end of each cycle, the four-carbon oxaloacetate has been regenerated, and the cycle continues. Because two acetyl CoA molecules are produced from each glucose molecule, two cycles are required per glucose molecule. After two turns of the cycle, the original glucose has lost all its carbons and may be regarded as having been completely consumed. To summarize, the citric acid cycle yields four CO2, six NADH, two FADH2, and two ATPs per glucose molecule. At this point in aerobic respiration, only four molecules of ATP have been formed per glucose by substrate-level phosphorylation: two during glycolysis and two during the citric acid cycle. Most of the energy of the original glucose molecule is in the form of highenergy electrons in NADH and FADH2. Their energy will be used to synthesize additional ATP through the electron transport chain and chemiosmosis.

The electron transport chain is coupled to ATP synthesis Let us consider the fate of all the electrons removed from a molecule of glucose during glycolysis, acetyl CoA formation, and the citric acid cycle. Recall that these electrons were transferred as part of hydrogen atoms to the acceptors NAD+ and FAD, forming NADH and FADH2. These reduced compounds now enter the electron transport chain, where the high-energy electrons of their hydrogen atoms are shuttled from one acceptor to another. As the electrons are passed along in a series of exergonic redox reactions, some of their energy is used to drive the synthesis of ATP, which is an endergonic process. Because ATP synthesis (by phosphorylation of ADP) is coupled to the redox reactions in the electron transport chain, the entire process is known as oxidative phosphorylation. The electron transport chain transfers electrons from NADH and FADH2 to oxygen The electron transport chain is a series of electron carriers embedded in the inner mitochondrial membrane of eukaryotes and in the plasma membrane of aerobic prokaryotes. Like NADH and FADH2, each carrier exists in an oxidized form or a reduced form. Electrons pass down the electron transport chain in a series of redox reactions that works much like a bucket brigade, the old-time chain of people who passed buckets of water from a stream to one another, to a building that was on fire. In the electron transport chain, each acceptor molecule becomes alternately reduced as it accepts electrons and oxidized as it gives them up. The electrons entering the electron transport chain have a relatively high energy content. They lose some of their energy at each step as they pass along the chain of electron carriers (just as some of the water spills out of the bucket as it is passed from one person to another).

FIGURE 8-4 A detailed look at glycolysis

CH2OH O

H

A specific enzyme catalyzes each of the reactions in glycolysis. Note the net yield of two ATP molecules and two NADH molecules. (The black wavy lines indicate bonds that permit the phosphates to be readily transferred to other molecules; in this case, ADP.)

H OH

Energy investment phase and splitting of glucose Two ATPs invested per glucose

H

H

HO

OH H

OH

Glucose ATP

Hexokinase

ADP CH2O

P O

H H OH

1

H

H

HO

OH H

OH

Glucose-6-phosphate

Glycolysis begins with preparation reaction in which glucose receives phosphate group from ATP molecule. ATP serves as source of both phosphate and energy needed to attach phosphate to glucose molecule. (Once ATP is spent, it becomes ADP and joins ADP pool of cell until turned into ATP again.) Phosphorylated glucose is known as glucose-6-phosphate. (Note phosphate attached to its carbon atom 6.) Phosphorylation of glucose makes it more chemically reactive.

Phosphoglucoisomerase

CH2O

P O

CH2OH

2

HO

H

OH

H HO

Glucose-6-phosphate undergoes another preparation reaction, rearrangement of its hydrogen and oxygen atoms. In this reaction glucose-6-phosphate is converted to its isomer, fructose-6-phosphate.

H

Fructose-6-phosphate ATP

Phosphofructokinase

ADP P

O

O

CH2

CH2

O

P

Next, another ATP donates phosphate to molecule, forming fructose-1,6-bisphosphate. So far, two ATP molecules have been invested in process without any being produced. Phosphate groups are now bound at carbons 1 and 6, and molecule is ready to be split.

3

H

HO OH

H HO

H

Fructose-1,6-bisphosphate Aldolase

P

O

H

CH2 C

C

O

4

Fructose-1,6-bisphosphate is then split into two 3-carbon sugars, glyceraldehyde-3phosphate (G3P) and dihydroxyacetone phosphate.

5

Dihydroxyacetone phosphate is enzymatically converted to its isomer, glyceraldehyde-3phosphate, for further metabolism in glycolysis.

O

Isomerase CH2OH

CHOH CH2

Dihydroxyacetone phosphate

O

P

Glyceraldehyde3-phosphate (G3P)

Two glyceraldehyde-3-phosphate (G3P) from bottom of previous page 2 NAD

Energy capture phase Four ATPs and two NADH produced per glucose

+

Glyceraldehyde-3-phosphate dehydrogenase

2 NADH

Pi O C

H

C

˜

P

6

OH

H 2C

P

O

Each glyceraldehyde-3-phosphate undergoes dehydrogenation with NAD+ as hydrogen acceptor. Product of this very exergonic reaction is phosphoglycerate, which reacts with inorganic phosphate present in cytosol to yield 1,3-bisphosphoglycerate.

Two 1,3-bisphosphoglycerate 2 ADP Phosphoglycerokinase 2 ATP O –

C

O

7

HC

OH

H 2C

O

P

One of phosphates of 1,3-bisphosphoglycerate reacts with ADP to form ATP. This transfer of phosphate from phosphorylated intermediate to ATP is referred to as substrate-level phosphorylation.

Two 3-phosphoglycerate Phosphoglyceromutase O O–

C

P

HC

O

H 2C

OH

8

3-phosphoglycerate is rearranged to 2-phosphoglycerate by enzymatic shift of position of phosphate group. This is a preparation reaction.

9

Next, molecule of water is removed, which results in formation of double bond. The product, phosphoenolpyruvate (PEP), has phosphate group attached by an unstable bond (wavy line).

Two 2-phosphoglycerate Enolase

2 H 2O

O C

O–

C

O

CH2

˜

P

Two phosphoenolpyruvate 2 ADP Pyruvate kinase 2 ATP O C

O–

C

O

CH3 Two pyruvate

10

Each of two PEP molecules transfers its phosphate group to ADP to yield ATP and pyruvate. This is substrate-level phosphorylation reaction.

Glycolysis

Formation of acetyl coenzyme A

Citric acid cycle

Glycolysis

Electron transport and chemiosmosis

Glucose

Glucose

Pyruvate

Pyruvate

2 ATP

2 ATP

Formation of acetyl coenzyme A

2 ATP

32 ATP

Acetyl coenzyme A

O H3C

Citric acid cycle

Electron transport and chemiosmosis

2 ATP

32 ATP

Coenzyme A

O C

C

O–

Carbon dioxide

CO2

NAD

NADH NAD+

Coenzyme A

NAD+

C IT R I C ACID CY C L E

H2O

Pyruvate +

Citrate

Oxaloacetate

NADH CO2

FADH2

NADH

5-carbon compound

FAD

NAD+

O H3C

˜

GTP

C S

CoA

NADH GDP

ADP

4-carbon compound

CO2

ATP

Acetyl coenzyme A

FIGURE 8-6 Animated Overview of the citric acid cycle FIGURE 8-5 The formation of acetyl CoA This series of reactions is catalyzed by the multienzyme complex pyruvate dehydrogenase. Pyruvate, a three-carbon molecule that is the end product of glycolysis, enters the mitochondrion and undergoes oxidative decarboxylation. First, the carboxyl group is split off as carbon dioxide. Then, the remaining two-carbon fragment is oxidized, and its electrons are transferred to NAD+. Finally, the oxidized two-carbon group, an acetyl group, is attached to coenzyme A. CoA has a sulfur atom that forms a bond, shown as a black wavy line, with the acetyl group. When this bond is broken, the acetyl group can be readily transferred to another molecule.

Members of the electron transport chain include the flavoprotein flavin mononucleotide (FMN), the lipid ubiquinone (also called coenzyme Q or CoQ), several iron–sulfur proteins, and a group of closely related iron-containing proteins called cytochromes (FIG. 8-8). Each electron carrier has a different mechanism for accepting and passing electrons. As cytochromes accept and donate electrons, for example, the charge on the iron atom, which is the electron carrier portion of the cytochromes, alternates between Fe2+ (reduced) and Fe3+ (oxidized). Scientists have extracted and purified the electron transport chain from the inner mitochondrial membrane as four large, distinct protein complexes, or groups, of acceptors. Complex I (NADH–ubiquinone oxidoreductase) accepts electrons from NADH

For every glucose, two acetyl groups enter the citric acid cycle (top). Each two-carbon acetyl group combines with a four-carbon compound, oxaloacetate, to form the six-carbon compound citrate. Two CO2 molecules are removed, and energy is captured as one ATP, three NADH, and one FADH2 per acetyl group (or two ATPs, six NADH, and two FADH2 per glucose molecule).

molecules that were produced during glycolysis, the formation of acetyl CoA, and the citric acid cycle. Complex II (succinate– ubiquinone reductase) accepts electrons from FADH2 molecules that were produced during the citric acid cycle. Complexes I and II both produce the same product, reduced ubiquinone, which is the substrate of complex III (ubiquinone–cytochrome c oxidoreductase). That is, complex III accepts electrons from reduced ubiquinone and passes them on to cytochrome c. Complex IV (cytochrome c oxidase) accepts electrons from cytochrome c and uses these electrons to reduce molecular oxygen, forming water in the process. The electrons simultaneously unite with protons from the surrounding medium to form hydrogen, and the chemical reaction between hydrogen and oxygen produces water. Because oxygen is the final electron acceptor in the electron transport chain, organisms that respire aerobically require oxygen. What happens when cells that are strict aerobes are deprived of oxygen? The last cytochrome in the chain retains its electrons when no oxygen is available to accept them. When that occurs, each

FIGURE 8-7 A detailed look at the citric acid cycle

COO–

8

Malate is dehydrogenated, forming oxaloacetate. Two hydrogens removed are transferred to NAD+. Oxaloacetate can now combine with another molecule of acetyl coenzyme A, beginning new cycle.

Unstable bond attaching acetyl group to coenzyme A breaks. 2-carbon acetyl group becomes attached to 4-carbon oxaloacetate molecule, forming citrate, 6-carbon molecule with three carboxyl groups. Coenzyme A is free to combine with another 2-carbon group and repeat process.

1

Begin with step 1, in the upper right corner, where acetyl coenzyme A attaches to oxaloacetate. Follow the steps in the citric acid cycle to see that the entry of a two-carbon acetyl group is balanced by the release of two molecules of CO2. Electrons are transferred to NAD+ or FAD, yielding NADH and FADH2, respectively, and ATP is formed by substratelevel phosphorylation.

H

C

O

C

H

Glucose

Acetyl coenzyme A

COO–

Malate dehydrogenase

Oxaloacetate COO– H

C

OH

H

C

H

Citrate synthase Coenzyme A

NADH

COO–

NAD+

COO–

Malate 7

With addition of water, fumarate is converted to malate.

Fatty acids

H

C

H

HO

C

COO–

H

C

H

COO–

Citrate H2O

Fumarase H2O

–

COO H

COO

Aconitase

H2O

C C

COO–

CITRIC ACID CYCLE

H –

Fumarate

HO

C

H

H

C

COO–

H

C COO

FADH2

6

Succinate is oxidized when two of its hydrogens are transferred to FAD, forming FADH2. Resulting compound is fumarate.

NAD+

Succinate dehydrogenase

FAD

C

H

H

C

H

COO–

NADH

COO –

NAD+ Coenzyme A

NADH

Succinate

GDP ATP

C

O

H

C

H

H

C

H

CO2

3

Isocitrate undergoes dehydrogenation and decarboxylation to yield 5-carbon compound α-ketoglutarate.

COO–

α-Ketoglutarate

Coenzyme A

ADP

H –

Isocitrate dehydrogenase

COO– H

Atoms of citrate are rearranged by two preparation reactions: first, molecule of water is removed, and then molecule of water is added. Through these reactions citrate is converted to its isomer, isocitrate.

Isocitrate

GTP Succinyl CoA synthetase

α -Ketoglutarate dehydrogenase

COO– CH2

4

CH2

5

2

Succinyl coenzyme A is converted to succinate, and substrate-level phosphorylation takes place. Bond attaching coenzyme A to succinate (~S) is unstable. Breakdown of succinyl coenzyme A is coupled to phosphorylation of GDP to form GTP (compound similar to ATP). GTP transfers its phosphate to ADP, yielding ATP.

C O

˜

S

CoA

Succinyl coenzyme A

CO2

Next α-ketoglutarate undergoes decarboxylation and dehydrogenation to form 4-carbon compound succinyl coenzyme A. Reaction is catalyzed by multienzyme complex similar to complex that catalyzes conversion of pyruvate to acetyl coenzyme A.

KEY POINT

Electron carriers in the mitochondrial inner membrane transfer electrons from NADH and FADH2 to oxygen.

Cytosol

Outer mitochondrial membrane

Intermembrane space

Inner mitochondrial membrane

Complex I: NADH–ubiquinone oxidoreductase

Complex II: Succinate– ubiquinone reductase

Matrix of mitochondrion

Complex III: Ubiquinone– cytochrome c oxidoreductase

Complex IV: Cytochrome c oxidase

FADH2 FAD

NAD

2 H

+

H2O 1 O 2 2

+

NADH

FIGURE 8-8 Animated An overview of the electron transport chain Electrons fall to successively lower energy levels as they are passed along the four complexes of the electron transport chain located in the inner mitochondrial membrane. (The orange arrows indicate the pathway of electrons.) The carriers within each complex become alternately

acceptor molecule in the chain retains its electrons (each remains in its reduced state), and the entire chain is blocked all the way back to NADH. Because oxidative phosphorylation is coupled to electron transport, no additional ATP is produced by way of the electron transport chain. Most cells of multicellular organisms cannot live long without oxygen because the small amount of ATP they produce by glycolysis alone is insufficient to sustain life processes. Lack of oxygen is not the only factor that interferes with the electron transport chain. Some poisons, including cyanide, inhibit the normal activity of the cytochromes. Cyanide binds tightly to the iron in the last cytochrome in the electron transport chain, making it unable to transport electrons to oxygen. This blocks the

reduced and oxidized as they accept and donate electrons. The terminal acceptor is oxygen; one of the two atoms of an oxygen molecule (written as –21 O2) accepts 2 electrons, which are added to 2 protons from the surrounding medium to produce water.

further passage of electrons through the chain, and ATP production ceases. Although the flow of electrons in electron transport is usually tightly coupled to the production of ATP, some organisms uncouple the two processes to produce heat (see Inquiring About: Electron Transport and Heat). The chemiosmotic model explains the coupling of ATP synthesis to electron transport in aerobic respiration For decades, scientists were aware that oxidative phosphorylation occurs in mitochondria, and many experiments had shown that the transfer of 2 electrons from each NADH to oxygen (via the

In q u ir in g A b o u t E L E C T RO N T R A N S P O RT A N D H E AT and wet woodlands and generally flowers during February and March when the ground is still covered with snow (see figure). Its uncoupled mitochondria generate large amounts of heat, enabling the plant to melt the snow and attract insect pollinators by vaporizing certain odiferous molecules into the surrounding air. The flower temperature of skunk cabbage is 15° to 22°C (59° to 72°F) when the air surrounding it is −15° to 10°C (5° to 50°F). Skunk cabbage flowers maintain this temperature for two weeks or more. Other plants, such as split leaf philodendron (Philodendron selloum) and sacred lotus (Nelumbo nucifera), also generate heat when they bloom and maintain their temperatures within precise limits. Some plants generate as much or more heat per gram of tissue than animals in flight, which have long been considered the greatest heat producers in the living world. The European plant lords-and-ladies (Arum maculatum), for example, produces 0.4 J (0.1 cal) of heat per second per gram of tissue, whereas a hummingbird in flight

electron transport chain) usually results in the production of up to three ATP molecules. However, for a long time, the connection between ATP synthesis and electron transport remained a mystery. In 1961, Peter Mitchell, a British biochemist, proposed the chemiosmotic model, which was based on his experiments and on theoretical considerations. One type of experiment involved using bacteria as a model system (FIG. 8-9). Because the respiratory electron transport chain is located in the plasma membrane of an aerobic bacterial cell, the bacterial plasma membrane can be considered comparable to the inner mitochondrial membrane. Mitchell demonstrated that if bacterial cells were placed in an acidic environment (that is, an environment with a high hydrogen ion, or proton, concentration), the cells synthesized ATP even if electron transport was not taking place. On the basis of these and other experiments, Mitchell proposed that electron transport and ATP synthesis are coupled by means of a proton gradient across the inner mitochondrial membrane in eukaryotes (or across the plasma membrane in bacteria). His model was so radical that it was not immediately accepted, but by 1978, so much evidence had accumulated in support of chemiosmosis that Peter Mitchell was awarded a Nobel Prize in Chemistry. The electron transport chain establishes the proton gradient; some of the energy released as electrons pass down the electron transport chain is used to move protons (H+) across a membrane. In eukaryotes the protons are moved across the inner mitochondrial membrane into the intermembrane space (FIG. 8-10). Hence, the inner mitochondrial

Leonard Lee Rue III/Animals Animals

What is the source of our body heat? Essentially, it is a byproduct of various exergonic reactions, especially those involving the electron transport chains in our mitochondria. Some cold-adapted animals, hibernating animals, and newborn animals produce unusually large amounts of heat by uncoupling electron transport from ATP production. These animals have adipose tissue (tissue in which fat is stored) that is brown. The brown color comes from the large number of mitochondria found in the brown adipose tissue cells. The inner mitochondrial membranes of these mitochondria contain an uncoupling protein that produces a passive proton channel through which protons flow into the mitochondrial matrix. As a consequence, most of the energy of glucose is converted to heat rather than to chemical energy in ATP. Certain plants, which are not generally considered “warm” organisms, also have the ability to produce large amounts of heat. Skunk cabbage (Symplocarpus foetidus), for example, lives in North American swamps

Skunk cabbage (Symplocarpus foetidus) This plant not only produces a significant amount of heat when it flowers but also regulates its temperature within a specific range.

produces 0.24 J (0.06 cal) per second per gram of tissue.

KEY EXPERIMENT QUESTION: What is the mechanism of oxidative phosphorylation? HYPOTHESIS: Peter Mitchell proposed that the cell uses energy released during electron transport to create a proton gradient across a membrane. The potential energy inherent in that gradient then drives the synthesis of ATP. EXPERIMENT: Aerobic bacteria were placed in an acid (high H+ concentration) environment, thus creating a proton gradient across the plasma membrane. This was done under conditions in which no electron transport was occurring.

+

H

H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+

H+

H+

Bacterial cytoplasm (low acid)

H+ H+

H+

H+ H+ H+

H+

H+

H+

H+

H+

H+

ATP

H+

Synthesized

H+

H+ +

+

H

H+

+

H

+

H

+

H

+

H

+

H

+

H

+

H

+

H

H

+

+

H

+

H

+

H

H

+

+

H

H+

H

Plasma membrane

Acidic environment

RESULTS AND CONCLUSION: The bacteria synthesized ATP in the absence of aerobic respiration. These results supported Mitchell’s view that a proton gradient across a membrane is an essential link in the conversion of the electrical energy of the electron transport chain to chemical energy in ATP.

FIGURE 8-9 Evidence for chemiosmosis

force of protons moving through the enzyme complex. The rotation apparently alters the conformation of the catalytic subunits in a way that drives ATP synthesis. Chemiosmosis is a fundamental mechanism of energy coupling in cells; it allows exergonic redox reactions to drive the endergonic reaction in which ATP is produced by phosphorylating ADP. In photosynthesis (discussed in Chapter 9), ATP is produced by a comparable process.

H+

H+

H+

Outer mitochondrial membrane

H+ H+

H+

H+

H+ + H+ H+ H

H+

H+

H+

H+

H+

H+

H+ H+

H+

H+

Intermembrane space—low pH

H+

H+

H+

H+ H+

H+ H+

H+

H+

Inner mitochondrial membrane

H+

H+

H+

Cytosol

H+

Matrix—higher pH

H+ H+ H+ H+

H+

FIGURE 8-10 The accumulation of protons (H+) within the intermembrane space As electrons move down the electron transport chain, the electron transport complexes move protons (H+) from the matrix to the intermembrane space, creating a proton gradient. The high concentration of H+ in the intermembrane space lowers the pH.

membrane separates a space with a higher concentration of protons (the intermembrane space) from a space with a lower concentration of protons (the mitochondrial matrix). Protons are moved across the inner mitochondrial membrane by three of the four electron transport complexes (complexes I, III, and IV) (FIG. 8-11a). Like water behind a dam, the resulting proton gradient is a form of potential energy that can be harnessed to provide the energy for ATP synthesis. Diffusion of protons from the intermembrane space, where they are highly concentrated, through the inner mitochondrial membrane to the matrix of the mitochondrion is limited to specific channels formed by a fifth enzyme complex, ATP synthase, a transmembrane protein. Portions of these complexes project from the inner surface of the membrane (the surface that faces the matrix) and are visible by electron microscopy (FIG. 8-11b). Diffusion of the protons down their gradient, through the ATP synthase complex, is exergonic because the entropy of the system increases. This exergonic process provides the energy for ATP production, although the exact mechanism by which ATP synthase catalyzes the phosphorylation of ADP is still not completely understood. In 1997, Paul Boyer of the University of California at Los Angeles and John Walker of the Medical Research Council Laboratory of Molecular Biology, Cambridge, England, shared the Nobel Prize in Chemistry for the discovery that ATP synthase functions in an unusual way. Experimental evidence strongly suggests that ATP synthase acts like a highly efficient molecular motor: During the production of ATP from ADP and inorganic phosphate, a central structure of ATP synthase rotates, possibly in response to the

Aerobic respiration of one glucose yields a maximum of 36 to 38 ATPs Let us now review where biologically useful energy is captured in aerobic respiration and calculate the total energy yield from the complete oxidation of glucose. FIGURE 8-12 summarizes the arithmetic involved. 1. In glycolysis, glucose is activated by the addition of phosphates from 2 ATP molecules and converted ultimately to 2 pyruvates + 2 NADH + 4 ATPs, yielding a net profit of 2 ATPs. 2. The 2 pyruvates are metabolized to 2 acetyl CoA + 2 CO2 + 2 NADH. 3. In the citric acid cycle, the 2 acetyl CoA molecules are metabolized to 4 CO2 + 6 NADH + 2 FADH2 + 2 ATPs. Because the oxidation of NADH in the electron transport chain yields up to 3 ATPs per molecule, the total of 10 NADH molecules can yield up to 30 ATPs. The 2 NADH molecules from glycolysis, however, yield either 2 or 3 ATPs each. The reason is that certain types of eukaryotic cells must expend energy to shuttle the NADH produced by glycolysis across the mitochondrial membrane (to be discussed shortly). Prokaryotic cells lack mitochondria; hence, they have no need to shuttle NADH molecules. For this reason, bacteria are able to generate 3 ATPs for every NADH, even those produced during glycolysis. Thus, the maximum number of ATPs formed using the energy from NADH is 28 to 30. The oxidation of FADH2 yields 2 ATPs per molecule (recall that electrons from FADH2 enter the electron transport chain at a different location than those from NADH), so the 2 FADH2 molecules produced in the citric acid cycle yield 4 ATPs. 4. Summing all the ATPs (2 from glycolysis, 2 from the citric acid cycle, and 32 to 34 from electron transport and chemiosmosis), you can see that the complete aerobic metabolism of one molecule of glucose yields a maximum of 36 to 38 ATPs. Most ATP is generated by oxidative phosphorylation, which involves the electron transport chain and chemiosmosis. Only 4 ATPs are formed by substrate-level phosphorylation in glycolysis and the citric acid cycle. We can analyze the efficiency of the overall process of aerobic respiration by comparing the free energy captured as ATP to the total free energy in a glucose molecule. Recall from Chapter 6 that although heat energy cannot power biological reactions, it is convenient to measure energy as heat. This is done through the use of a calorimeter, an instrument that measures the heat of a reaction. A sample is placed in a compartment surrounded by a chamber of water. As the sample burns (becomes oxidized), the temperature of the water rises, providing a measure of the heat released during the reaction.

KEY POINT

The electron transport chain forms a concentration gradient for H+, which diffuses through ATP synthase complexes, producing ATP. Glycolysis

Formation of acetyl coenzyme A

Citric acid cycle

Electron transport and chemiosmosis

2 ATP

32 ATP

Glucose

Pyruvate

2 ATP

Cytosol Outer mitochondrial membrane Intermembrane space

H

H+

H+

H+

H+ +

H+

H+

H+

H

H+

H+ H+

H+

H+ H

H

Complex II

Complex I

Matrix of mitochondrion

H+

+

+

Inner mitochondrial membrane

H+

H+

H+

+

H+

H+

H+

Complex III

Complex V: ATP synthase

Complex IV

FADH2 FAD NAD+

1 O 2 2

NADH H+

H+

H2O

2 H+

H+

H+ ADP + Pi

R. Bhatnagar/Visuals Unlimited, Inc.

(a) The electron transport chain in the inner mitochondrial membrane includes three proton pumps that are located in three of the four electron transport complexes. (The orange arrows indicate the pathway of electrons; and the black arrows, the pathway of protons.) The energy released during electron transport is used to transport protons (H+ ) from the mitochondrial matrix to the intermembrane space, where a high concentration of protons accumulates. The protons cannot diffuse back into the matrix except through special channels in ATP synthase in the inner membrane. The flow of the protons through ATP synthase provides the energy for generating ATP from ADP and inorganic phosphate (Pi ). In the process, the inner part of ATP synthase rotates (thick red arrows) like a motor.

ATP

Projections of ATP synthase

250 nm

(b) This TEM shows hundreds of projections of ATP synthase complexes along the surface of the inner mitochondrial membrane.

FIGURE 8-11 A detailed look at electron transport and chemiosmosis

KEY POINT Substrate-level phosphorylation

Oxidative phosphorylation

Glycolysis Glucose

2

are transferred to the electron transport chain in the inner mitochondrial membrane, and up to three molecules of ATP are produced per pair of electrons. In skeletal muscle, brain, and some other types of cells, another type of shuttle operates. Because this shuttle requires more energy than the shuttle in liver, kidney, and heart cells, the electrons are at a lower energy level when they enter the electron transport chain. They are accepted by ubiquinone rather than by NAD+ and so generate a maximum of 2 ATP molecules per pair of electrons. For this reason, the number of ATPs produced by aerobic respiration of 1 molecule of glucose in skeletal muscle cells is 36 rather than 38.

Most ATP is produced by electron transport and chemiosmosis (oxidative phosphorylation)

ATP

2 NADH

4–6

ATP

2 NADH

6

ATP

Pyruvate

Acetyl coenzyme A

Cells regulate aerobic respiration

2

4

ATP

ATP

Total ATP from substratelevel phosphorylation

Citric acid cycle

6 NADH

18

ATP

2 FADH2

4

ATP

32 – 34

ATP

Total ATP from oxidative phosphorylation

FIGURE 8-12 Energy yield from the complete oxidation of glucose by aerobic respiration

When 1 mol of glucose is burned in a calorimeter, some 686 kcal (2870 kJ) are released as heat. The free energy temporarily held in the phosphate bonds of ATP is about 7.6 kcal (31.8 kJ) per mole. When 36 to 38 ATPs are generated during the aerobic respiration of glucose, the free energy trapped in ATP amounts to 7.6 kcal/mol × 36, or about 274 kcal (1146 kJ) per mole. Thus, the efficiency of aerobic respiration is 274/686, or about 40%. (By comparison, a steam power plant has an efficiency of 35% to 36% in converting its fuel energy into electricity.) The remainder of the energy in the glucose is released as heat. Mitochondrial shuttle systems harvest the electrons of NADH produced in the cytosol The inner mitochondrial membrane is not permeable to NADH, which is a large molecule. Therefore, the NADH molecules produced in the cytosol during glycolysis cannot diffuse into the mitochondria to transfer their electrons to the electron transport chain. Unlike ATP and ADP, NADH does not have a carrier protein to transport it across the membrane. Instead, several systems have evolved to transfer just the electrons of NADH, not the NADH molecules themselves, into the mitochondria. In liver, kidney, and heart cells, a special shuttle system transfers the electrons from NADH through the inner mitochondrial membrane to an NAD+ molecule in the matrix. These electrons

Aerobic respiration requires a steady input of fuel molecules and oxygen. Under normal conditions these materials are adequately provided and do not affect the rate of respiration. Instead, the rate of aerobic respiration is regulated by how much ADP and phosphate are available, with ATP synthesis continuing until most of the ADP has been converted to ATP. At this point oxidative phosphorylation slows considerably, which in turn slows down the citric acid cycle. Glycolysis is partly controlled by feedback regulation (see Fig. 7-15 for an illustration of feedback regulation) exerted on the enzyme phosphofructokinase, which catalyzes an early reaction of glycolysis (see Fig. 8-4). The active site of phosphofructokinase binds ATP and fructose6-phosphate. However, the enzyme has two allosteric sites: an inhibitor site to which ATP binds when present at very high levels, and an activator site to which AMP (adenosine monophosphate, a molecule formed when two phosphates are removed from ATP) binds. Therefore, this enzyme is inactivated when ATP levels are high and activated when they are low. Respiration proceeds when the enzyme becomes activated, thus generating more ATP.

Review ■

■

■ ■

How much ATP is made available to the cell from a single glucose molecule by the operation of (1) glycolysis, (2) the formation of acetyl CoA, (3) the citric acid cycle, and (4) the electron transport chain and chemiosmosis? Why is each of the following essential to chemiosmotic ATP synthesis: (1) electron transport chain, (2) proton gradient, and (3) ATP synthase complex? What are the roles of NAD+, FAD, and oxygen in aerobic respiration? What are some of the ways aerobic respiration is controlled?

8.3 ENERGY YIELD OF NUTRIENTS OTHER THAN GLUCOSE ■ ■ LEARNING OBJECTIVE 7

Summarize how the products of protein and lipid catabolism enter the same metabolic pathway that oxidizes glucose.

Many organisms, including humans, depend on nutrients other than glucose as a source of energy. In fact, you usually obtain more of your energy by oxidizing fatty acids than by oxidizing glucose. Amino acids derived from protein digestion are also used as fuel molecules. Such nutrients are transformed into one of the metabolic intermediates that are fed into glycolysis or the citric acid cycle (FIG. 8-13). Amino acids are metabolized by reactions in which the amino group (¬NH2) is first removed, a process called deamination. In mammals and some other animals, the amino group is converted to urea (see Fig. 48-1 for the biochemical pathway) and excreted, but the carbon chain is metabolized and eventually is used as a reactant in one of the steps of aerobic respiration. The amino acid alanine, for example, undergoes deamination to become pyruvate, the amino acid glutamate is converted to a-ketoglutarate, and the amino acid aspartate yields oxaloacetate. Pyruvate enters aerobic respiration as the end product of glycolysis, and a-ketoglutarate and oxaloacetate both enter aerobic respiration as intermediates in the citric acid cycle. Ultimately, the carbon chains of all the amino acids are metabolized in this way.

Each gram of lipid in the diet contains 9 kcal (38 kJ), more than twice as much energy as 1 g of glucose or amino acids, which have about 4 kcal (17 kJ) per gram. Lipids are rich in energy because they are highly reduced; that is, they have many hydrogen atoms and few oxygen atoms. When completely oxidized in aerobic respiration, a molecule of a six-carbon fatty acid generates up to 44 ATPs (compared with 36 to 38 ATPs for a molecule of glucose, which also has six carbons). Both the glycerol and fatty acid components of a triacylglycerol (see Figure 3-12 for structures) are used as fuel; phosphate is added to glycerol, converting it to G3P or another compound that enters glycolysis. Fatty acids are oxidized and split enzymatically into two-carbon acetyl groups that are bound to coenzyme A; that is, fatty acids are converted to acetyl CoA. This process, which occurs in the mitochondrial matrix, is called b-oxidation (beta-oxidation). Acetyl CoA molecules formed by b-oxidation enter the citric acid cycle.

Review ■ ■

PROTEINS

CARBOHYDRATES

FATS ■

Amino acids

Glycolysis

Glycerol

Fatty acids

Glucose

8.4 ANAEROBIC RESPIRATION AND FERMENTATION

G3P

■ ■ LEARNING OBJECTIVE

Pyruvate

8

CO2 Acetyl coenzyme A

Electron transport and chemiosmosis

NH3

H2O

Compare and contrast anaerobic respiration and fermentation. Include the mechanism of ATP formation, the final electron acceptor, and the end products.

Anaerobic respiration, which does not use oxygen as the final electron acceptor, is performed by some prokaryotes that live in anaerobic environments, such as waterlogged soil, stagnant ponds, and animal intestines. As in aerobic respiration, electrons are transferred in anaerobic respiration from glucose to NADH; they then pass down an electron transport chain that is coupled to ATP synthesis by chemiosmosis. However, an inorganic substance such as nitrate (NO3−) or sulfate (SO42−) replaces molecular oxygen as the terminal electron acceptor. The end products of this type of anaerobic respiration are carbon dioxide, one or more reduced inorganic substances, and ATP. The following equation summarizes one representative type of anaerobic respiration, which is part of the biogeochemical cycle known as the nitrogen cycle (discussed in Chapter 55).

Citric acid cycle

End products:

How can a person obtain energy from a low-carbohydrate diet? What process must occur before amino acids enter the aerobic respiratory pathway? Where do fatty acids enter the aerobic respiratory pathway?

CO2

FIGURE 8-13 Animated Energy from proteins, carbohydrates, and fats Products of the catabolism of proteins, carbohydrates, and fats enter glycolysis or the citric acid cycle at various points. This diagram is greatly simplified and illustrates only a few of the principal catabolic pathways.

C 6H12 O 6 + 12 KNO3 ¡ Potassium nitrate

6 CO2 + 6 H2O + 12 KNO2 + energy Potassium (in the chemical nitrite bonds of ATP)

TABLE 8-2

A Comparison of Aerobic Respiration, Anaerobic Respiration, and Fermentation

Immediate fate of electrons in NADH Terminal electron acceptor of electron transport chain Reduced product(s) formed

Mechanism of ATP synthesis

Aerobic Respiration

Anaerobic Respiration

Fermentation

Transferred to electron transport chain O2

Transferred to electron transport chain

Transferred to organic molecule

Inorganic substances such as NO3− or SO42− Relatively reduced inorganic substances

No electron transport chain

Water

Oxidative phosphorylation/ chemiosmosis; also substratelevel phosphorylation

Oxidative phosphorylation/chemiosmosis; also substrate-level phosphorylation

Certain other bacteria, as well as some fungi, regularly use fermentation, an anaerobic pathway that does not involve an electron transport chain. During fermentation only two ATPs are formed per glucose (by substrate-level phosphorylation during glycolysis). One might expect that a cell that obtains energy from glycolysis would produce pyruvate, the end product of glycolysis. However, this cannot happen because every cell has a limited supply of NAD+, and NAD+ is required for glycolysis to continue. If virtually all NAD+ becomes reduced to NADH during glycolysis, then glycolysis stops and no more ATP is produced. In fermentation, NADH molecules transfer their hydrogen atoms to organic molecules, thus regenerating the NAD+ needed to keep glycolysis going. The resulting relatively reduced organic molecules (commonly, alcohol or lactate) tend to be toxic to the cells and are essentially waste products.

KEY POINT

Relatively reduced organic compounds (commonly, alcohol or lactate) Substrate-level phosphorylation only (during glycolysis)

TABLE 8-2 compares aerobic respiration, anaerobic respiration, and fermentation.

Alcohol fermentation and lactate fermentation are inefficient Yeasts are facultative anaerobes that carry out aerobic respiration when oxygen is available but switch to alcohol fermentation when deprived of oxygen (FIG. 8-14a). These eukaryotic, unicellular fungi have enzymes that decarboxylate pyruvate, releasing carbon dioxide and forming a two-carbon compound called acetaldehyde. NADH produced during glycolysis transfers hydrogen atoms to acetaldehyde, reducing it to ethyl alcohol (FIG. 8-14b). Alcohol fermentation is the basis for the production of beer, wine, and other alcoholic beverages. Yeast cells are also used in baking to

Fermentation regenerates NAD+ needed for glycosis.

Glycolysis

Glycolysis

Glucose

Glucose

2 NAD+

2 NADH

Dwight R. Kuhn

2 ATP

25 μm

2 NAD+

2 NADH

2 ATP 2 Pyruvate

2 Pyruvate

2 Ethyl alcohol

2 Lactate

CO2

(a) Yeast cells (b) Alcohol fermentation

(c) Lactate fermentation

FIGURE 8-14 Animated Fermentation (a) Light micrograph of live brewer’s yeast (Saccharomyces cerevisiae). Yeast cells have mitochondria and carry on aerobic respiration when O2 is present. In the absence of O2, yeasts carry on alcohol fermentation. (b, c) Glycolysis is the first part of fermentation pathways. In alcohol fermentation (b), CO2 is split off, and the two-carbon compound ethyl

alcohol is the end product. In lactate fermentation (c), the final product is the three-carbon compound lactate. In both alcohol and lactate fermentation, there is a net gain of only two ATPs per molecule of glucose. Note that the NAD+ used during glycolysis is regenerated during both alcohol fermentation and lactate fermentation.

produce the carbon dioxide that causes dough to rise; the alcohol evaporates during baking. Certain fungi and bacteria perform lactate (lactic acid) fermentation. In this alternative pathway, NADH produced during glycolysis transfers hydrogen atoms to pyruvate, reducing it to lactate (FIG. 8-14c). The ability of some bacteria to produce lactate is exploited by humans, who use these bacteria to make yogurt and to ferment cabbage for sauerkraut. Vertebrate muscle cells also produce lactate. Exercise can cause fatigue and muscle cramps possibly due to insufficient oxygen, the depletion of fuel molecules, and the accumulation of lactate during strenuous activity. This buildup of lactate occurs because muscle cells shift briefly to lactate fermentation if the amount of oxygen delivered to muscle cells is insufficient to support aerobic respiration. The shift is only temporary, however, and oxygen is required for sustained work. About 80% of the lactate is eventually exported to the liver, where it is used to regenerate more glucose for the muscle cells. The remaining 20% of the lactate is metabolized in muscle cells in the presence of oxygen. For this reason, you continue to breathe heavily after you have stopped exercising: the additional oxygen is needed to oxidize lactate, thereby restoring the muscle cells to their normal state. Although humans use lactate fermentation to produce ATP for only a few minutes, a few animals can live without oxygen for much longer periods. The red-eared slider, a freshwater turtle, remains underwater for as long as two weeks. During this time, it is relatively inactive and therefore does not expend a great deal of energy. It relies on lactate fermentation for ATP production.

■ ■

Both alcohol fermentation and lactate fermentation are highly inefficient because the fuel is only partially oxidized. Alcohol, the end product of fermentation by yeast cells, can be burned and is even used as automobile fuel; obviously, it contains a great deal of energy that the yeast cells cannot extract using anaerobic methods. Lactate, a three-carbon compound, contains even more energy than the two-carbon alcohol. In contrast, all available energy is removed during aerobic respiration because the fuel molecules become completely oxidized to CO2. A net profit of only 2 ATPs is produced by the fermentation of one molecule of glucose, compared with up to 36 to 38 ATPs when oxygen is available. The inefficiency of fermentation necessitates a large supply of fuel. To perform the same amount of work, a cell engaged in fermentation must consume up to 20 times as much glucose or other carbohydrate per second as a cell using aerobic respiration. For this reason, your skeletal muscle cells store large quantities of glucose in the form of glycogen, which enables them to metabolize anaerobically for short periods.

Review What is the fate of hydrogen atoms removed from glucose during glycolysis when oxygen is present in muscle cells? How does this compare to the fate of hydrogen atoms removed from glucose when the amount of available oxygen is insufficient to support aerobic respiration? Why is the ATP yield of fermentation only a tiny fraction of the yield from aerobic respiration?

■

■

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

8.1 (page 173)

■

1 Write a summary reaction for aerobic respiration that shows which reactant becomes oxidized and which becomes reduced. ■

Oxidation

C 6 H12 O6 ⫹ 6 O2

¡ 6 CO2 ⫹ 6 H2 O ⫹ energy Reduction

■

■

Aerobic respiration is a catabolic process in which a fuel molecule such as glucose is broken down to form carbon dioxide and water. It includes redox reactions that result in the transfer of electrons from glucose (which becomes oxidized) to oxygen (which becomes reduced). Energy released during aerobic respiration is used to produce up to 36 to 38 ATPs per molecule of glucose.

Interact with the citric acid cycle by clicking on the figure in CengageNOW. ■

8.2 (page 173) 2 List and give a brief overview of the four stages of aerobic respiration. ■

■

The chemical reactions of aerobic respiration occur in four stages: glycolysis, formation of acetyl CoA, the citric acid cycle, and the electron transport chain and chemiosmosis. During glycolysis, a molecule of glucose is degraded to two molecules of pyruvate. Two ATP molecules (net) are produced by substrate-level phosphorylation during glycolysis. Four hydrogen atoms are removed and used to produce two NADH.

During the formation of acetyl CoA, the two pyruvate molecules each lose a molecule of carbon dioxide, and the remaining acetyl groups each combine with coenzyme A, producing two molecules of acetyl CoA; one NADH is produced per pyruvate. Each acetyl CoA enters the citric acid cycle by combining with a four-carbon compound, oxaloacetate, to form citrate, a six-carbon compound. Two acetyl CoA molecules enter the cycle for every glucose molecule. For every two carbons that enter the cycle as part of an acetyl CoA molecule, two leave as carbon dioxide. For every acetyl CoA, hydrogen atoms are transferred to three NAD+ and one FAD; only one ATP is produced by substrate-level phosphorylation.

Hydrogen atoms (or their electrons) removed from fuel molecules are transferred from one electron acceptor to another down an electron transport chain located in the mitochondrial inner membrane; ultimately, these electrons reduce molecular oxygen, forming water. In oxidative phosphorylation, the redox reactions in the electron transport chain are coupled to synthesis of ATP through the mechanism of chemiosmosis. See the electron transport chain in action by clicking on the figure in CengageNOW.

3 Indicate where each stage of aerobic respiration takes place in a euSee the process of glycolysis unfold by clicking on the figure in CengageNOW.

karyotic cell.

Glycolysis

Formation of acetyl coenzyme A

Citric acid cycle

Electron transport and chemiosmosis

Glucose Mitochondrion Acetyl coenzyme A

Citric acid cycle

Electron transport and chemiosmosis

Pyruvate

2 ATP

Glycolysis occurs in the cytosol, and the remaining stages of aerobic respiration take place in the mitochondria. 4 Add up the energy captured (as ATP, NADH, and FADH2) in each stage of aerobic respiration. In glycolysis, each glucose molecule produces 2 NADH and 2 ATPs ■ (net). The conversion of 2 pyruvates to acetyl CoA results in the formation of 2 NADH. In the citric acid cycle, the 2 acetyl CoA molecules are metabolized to form 6 NADH, 2 FADH2, and 2 ATPs. To summarize, we have 4 ATPs, 10 NADH, and 2 FADH2. ■ When electrons donated by the 10 NADH and 2 FADH2 pass through the electron transport chain, 32 to 34 ATPs are produced by chemiosmosis. Therefore, each glucose molecule yields a total of up to 36 to 38 ATPs. 5 Define chemiosmosis and explain how a gradient of protons is established across the inner mitochondrial membrane. ■ In chemiosmosis, some of the energy of the electrons in the electron transport chain is used to pump protons across the inner mitochondrial membrane into the intermembrane space. This pumping establishes a proton gradient across the inner mitochondrial membrane. Protons (H+) accumulate within the intermembrane space, lowering the pH. 6 Describe the process by which the proton gradient drives ATP synthesis in chemiosmosis. ■ The diffusion of protons through channels formed by the enzyme ATP synthase, which extends through the inner mitochondrial membrane from the intermembrane space to the mitochondrial matrix, provides the energy to synthesize ATP. ■

2 ATP

■

■

■

■

32 ATP

In anaerobic respiration, electrons are transferred from fuel molecules to an electron transport chain that is coupled to ATP synthesis by chemiosmosis; the final electron acceptor is an inorganic substance such as nitrate or sulfate, not molecular oxygen. Fermentation is an anaerobic process that does not use an electron transport chain. There is a net gain of only two ATPs per glucose; these are produced by substrate-level phosphorylation during glycolysis. To maintain the supply of NAD+ essential for glycolysis, hydrogen atoms are transferred from NADH to an organic compound derived from the initial nutrient. Yeast cells carry out alcohol fermentation, in which ethyl alcohol and carbon dioxide are the final waste products. Certain fungi, prokaryotes, and animal cells carry out lactate (lactic acid) fermentation, in which hydrogen atoms are added to pyruvate to form lactate, a waste product. Learn more about fermentation by clicking on the figure in CengageNOW.

Summary Reactions for Aerobic Respiration Summary reaction for the complete oxidation of glucose: C6H12O6 + 6 O2 + 6 H2O ¡ 6 CO2 + 12 H2O + energy (36 to 38 ATP) Summary reaction for glycolysis: C6H12O6 + 2 ATP + 2 ADP + 2 Pi + 2 NAD+ ¡ 2 pyruvate + 4 ATP + 2 NADH + H2O

8.3 (page 186) 7 Summarize how the products of protein and lipid catabolism enter the same metabolic pathway that oxidizes glucose. ■ Amino acids undergo deamination, and their carbon skeletons are converted to metabolic intermediates of aerobic respiration. ■ Both the glycerol and fatty acid components of lipids are oxidized as fuel. Fatty acids are converted to acetyl CoA molecules by the process of b-oxidation.

8.4 (page 187) 8 Compare and contrast anaerobic respiration and fermentation. Include the mechanism of ATP formation, the final electron acceptor, and the end products.

Summary reaction for the conversion of pyruvate to acetyl CoA: 2 pyruvate + 2 coenzyme A + 2 NAD+ ¡ 2 acetyl CoA + 2 CO2 + 2 NADH Summary reaction for the citric acid cycle: 2 acetyl CoA + 6 NAD+ + 2 FAD + 2 ADP + 2 Pi + 2 H2O ¡ 4 CO2 + 6 NADH + 2 FADH2 + 2 ATP + 2 CoA

Summary reactions for the processing of the hydrogen atoms of NADH and FADH2 in the electron transport chain: NADH + 3 ADP + 3

Pi + –21 O2

Summary Reactions for Fermentation Summary reaction for lactate fermentation:

+

¡ NAD + 3 ATP + H2O

FADH2 + 2 ADP + 2 Pi + –21 O2 ¡ FAD+ + 2 ATP + H2O

C6H12O6 ¡ 2 lactate + energy (2 ATP) Summary reaction for alcohol fermentation: C6H12O6 ¡ 2 CO2 + 2 ethyl alcohol + energy (2 ATP)

T E S T YO U R U N D E R S TA N D I N G 1. A chemical process during which a substance gains electrons and energy is called (a) oxidation (b) oxidative phosphorylation (c) deamination (d) reduction (e) dehydrogenation 2. Which of the following is a correct ranking of molecules with respect to their energy value in glycolysis (note: > means “greater than”)? (a) two pyruvates > one glucose (b) one glucose > one fructose-1,6-bisphosphate (c) two glyceraldehyde-3-phosphates (G3P) > one glucose (d) two pyruvates > one fructose-1,6bisphosphate (e) two pyruvates > two glyceraldehyde-3phosphates (G3P) 3. The reactions of _____ take place within the cytosol of eukaryotic cells. (a) glycolysis (b) oxidation of pyruvate (c) the citric acid cycle (d) chemiosmosis (e) the electron transport chain

(note: > means “greater than”)? (a) ATP = NADH (b) NAD+ > NADH (c) FAD > FADH2 (d) NADH > ATP 10. A net profit of only 2 ATPs can be produced anaerobically from the _____ of one molecule of glucose, compared with a maximum of 38 ATPs produced in _____. (a) fermentation; anaerobic respiration (b) aerobic respiration; fermentation (c) aerobic respiration; anaerobic respiration (d) dehydrogenation; decarboxylation (e) fermentation; aerobic respiration 11. When deprived of oxygen, yeast cells obtain energy by fermentation, producing carbon dioxide, ATP, and (a) acetyl CoA (b) ethyl alcohol (c) lactate (d) pyruvate (e) citrate 12. Label the figure. Use Figure 8-10 to check your answers.

4. Before pyruvate enters the citric acid cycle, it is decarboxylated, oxidized, and combined with coenzyme A, forming acetyl CoA, carbon dioxide, and one molecule of (a) NADH (b) FADH2 (c) ATP (d) ADP (e) C6H12O6 5. In the first step of the citric acid cycle, acetyl CoA reacts with oxaloacetate to form (a) pyruvate (b) citrate (c) NADH (d) ATP (e) CO2 6. Which of the following is the major source of electrons that flow through the mitochondrial electron transport chain? (a) H2O (b) ATP (c) NADH (d) ATP synthase (e) coenzyme A 7. The “aerobic” part of aerobic cellular respiration occurs during (a) glycolysis (b) the conversion of pyruvate to acetyl CoA (c) the citric acid cycle (d) electron transport (e) all of the above are aerobic processes 8. Substrate-level phosphorylation (a) occurs through a chemiosmotic mechanism (b) accounts for most of the ATP formed during aerobic cellular respiration (c) occurs during the conversion of pyruvate to acetyl CoA (d) occurs during glycolysis and the citric acid cycle (e) requires high energy electrons from NADH 9. Which of the following is a correct ranking of molecules, according to their energy value in oxidative phosphorylation

H+

H+

H+

H+ H+

H+

H+

H+ H+

H+ + H+ H+ H

H+ H+

H+ H+

H+

H+ H+

H+

H+

H+

H+ H+ H+

H+

H+

H+

H+ H+

H+

H+

H+ H+ H+ H+

H+

H+

CRITICAL THINKING 1. How are the endergonic reactions of the first phase of glycolysis coupled to the hydrolysis of ATP, which is exergonic? How are the exergonic reactions of the second phase of glycolysis coupled to the endergonic synthesis of ATP and NADH?

5. EVOLUTION LINK. The reactions of glycolysis are identical in all organisms—prokaryotes, protists, fungi, plants, and animals— that obtain energy from glucose catabolism. What does this universality suggest about the evolution of glycolysis?

2. In what ways is the inner mitochondrial membrane essential to the coupling of electron transport and ATP synthesis? Could the membrane carry out its function if its lipid bilayer were readily permeable to hydrogen ions (protons)?

6. EVOLUTION LINK. Molecular oxygen is so reactive that it would not exist in Earth’s atmosphere today if it were not constantly replenished by organisms that release oxygen as a waste product of photosynthesis. What does that fact suggest about the evolution of aerobic respiration and oxygen-releasing photosynthetic processes?

3. Based on what you have learned in this chapter, explain why a schoolchild can run 17 miles per hour in a 100-yard dash, but a trained athlete can run only about 11.5 miles per hour in a 26-mile marathon. 4. When you lose weight, where does it go?

Additional questions are available in CengageNOW at www.cengage.com/ login.

9

Photosynthesis: Capturing Light Energy

© tbkmedia.de/Alamy

Photosynthesis. These trees use light energy to power the processes that incorporate CO2 into organic molecules.

KEY CONCEPTS

L

ook at all the living things that surround you—the trees, your pet goldfish, your own body. Most of that biomass is made up of carbon-based

9.1 Light energy powers photosynthesis, which is essen-

biological molecules. What is the ultimate source of all that carbon?

tial to plants and most life on Earth.

Surprising to some, the source is carbon dioxide from the air. Your cells

9.2 Photosynthesis occurs in chloroplasts and requires the

cannot take carbon dioxide from the air and incorporate it into organic

pigment chlorophyll.

molecules—but some plant cells can. They do this through photosynthe-

9.3 Photosynthesis is a redox process.

sis, the sequence of events by which light energy is converted into the

9.4 Light-dependent reactions convert light energy to

stored chemical energy of organic molecules. Photosynthesis is the first

the chemical energy of NADPH and ATP.

step in the flow of energy through most of the living world, capturing

9.5 Carbon fixation reactions incorporate CO2 into

the vast majority of the energy that living organisms use. Photosynthe-

organic molecules.

9.6 Most photosynthetic organisms are photoautotrophs. 9.7 Photosynthesis is important to plants and also other organisms.

sis not only sustains plants (see photograph) and other photosynthetic organisms such as algae and photosynthetic bacteria but also indirectly supports most nonphotosynthetic organisms such as animals, fungi, protozoa, and most bacteria. Each year photosynthetic organisms convert CO2 into billions of tons of organic molecules. These molecules have two important roles in both photosynthetic and nonphotosynthetic organisms: they are both the building blocks of cells and, as we saw in Chapter 8, a source of chemical energy that fuels the metabolic reactions that sustain almost all life. Photosynthesis also releases O2, which is essential to aerobic cellular respiration, the process by which plants, animals, and most other organisms convert this chemical energy to ATP to power cellular processes.

In this chapter we first examine how light energy is used in the

One wavelength

synthesis of ATP and other molecules that temporarily hold chemical energy but are unstable and cannot be stockpiled in the cell. We

Longer wavelength

then see how their energy powers the anabolic pathway by which a photosynthetic cell synthesizes stable organic molecules from the simple inorganic compounds CO2 and water. Finally, we explore the role of photosynthesis in plants and in Earth’s environment.

9.1 LIGHT AND PHOTOSYNTHESIS ■ ■ LEARNING OBJECTIVE 1

Describe the physical properties of light and explain the relationship between a wavelength of light and its energy.

Because most life on this planet depends on light, either directly or indirectly, it is important to understand the nature of light and its essential role in photosynthesis. Visible light represents a very small portion of a vast, continuous range of radiation called the electromagnetic spectrum (FIG. 9-1). All radiation in this spectrum travels as waves. A wavelength is the distance from one wave peak to the next. At one end of the electromagnetic spectrum are gamma rays, which have very short wavelengths measured in fractions of nanometers, or nm (1 nanometer equals 10–9 m, one-billionth of a meter). At the other end of the spectrum are radio waves, with wavelengths so long they can be measured in kilometers. The portion of the electromagnetic spectrum from 380 to 760 nm is called the visible spectrum because we humans can see it. The visible spectrum includes all the colors of the rainbow (FIG. 9-2); violet has the shortest wavelength, and red has the longest. Light is composed of small particles, or packets, of energy called photons. The energy of a photon is inversely proportional to its wavelength: shorter-wavelength light has more energy per photon than longer-wavelength light. Why does photosynthesis depend on light detectable by the human eye (visible light) rather than on some other wavelength of radiation? We can only speculate about the answer. Perhaps the reason is that radiation within the visible-light portion of the spectrum excites certain types of biological molecules, moving electrons into higher energy levels. Radiation with wavelengths longer than those of visible light does not have enough energy to excite these biological molecules. Radiation with wavelengths shorter than those of visible light is so energetic that it disrupts the bonds of many biological molecules. Thus, visible light has just the right amount of energy to cause the kinds of reversible changes in molecules that are useful in photosynthesis. When a molecule absorbs a photon of light energy, one of its electrons becomes energized, which means that the electron shifts from a lower-energy atomic orbital to a high-energy orbital that is more distant from the atomic nucleus. One of two things then happens to the energized electron, depending on the atom and its

760 nm

TV and radio waves

Red

700 nm

Microwaves Infrared Visible UV X-rays

Gamma rays

Orange Color spectrum of visible light

600 nm Yellow Green 500 nm Blue

Violet Electromagnetic spectrum

400 nm 380 nm

Shorter wavelength

FIGURE 9-1 Animated The electromagnetic spectrum Waves in the electromagnetic spectrum have similar properties but different wavelengths. Radio waves are the longest (and least energetic) waves, with wavelengths as long as 20 km. Gamma rays are the shortest (and most energetic) waves. Visible light represents a small fraction of the electromagnetic spectrum and consists of a mixture of wavelengths ranging from about 380 to 760 nm. The energy from visible light is used in photosynthesis.

surroundings (FIG. 9-3). The atom may return to its ground state, which is the condition in which all its electrons are in their normal, lowest-energy levels. When an electron returns to its ground state, its energy dissipates as heat or as an emission of light of a longer wavelength than the absorbed light; this emission of light is called fluorescence. Alternatively, the energized electron may leave the atom and be accepted by an electron acceptor molecule,

Sun Sunlight is a mixture of many wavelengths

FIGURE 9-2 Visible radiation emitted from the sun Electromagnetic radiation from the sun includes ultraviolet radiation and visible light of varying colors and wavelengths.

Photon

Photon is absorbed by an excitable electron that moves into a higher energy level.

Low energy level High energy level

Electron

Either

Or

Electron acceptor molecule The electron may return to ground level by emitting a less energetic photon.

The electron may be accepted by an electron acceptor molecule.

FIGURE 9-3 Interactions between light and atoms or molecules (Top) When a photon of light energy strikes an atom or a molecule of which the atom is a part, the energy of the photon may push an electron to an orbital farther from the nucleus (that is, into a higher energy level). (Lower left) If the electron returns to the lower, more stable energy level, the energy may be released as a less energetic, longer-wavelength photon, known as fluorescence (shown), or as heat. (Lower right) If the appropriate electron acceptors are available, the electron may leave the atom. During photosynthesis, an electron acceptor captures the energetic electron and passes it to a chain of acceptors.

which becomes reduced in the process; this is what occurs in photosynthesis. Now that you understand some of the properties of light, let us consider the organelles that use light for photosynthesis.

Review ■ ■ ■

Why does photosynthesis require visible light? Which color of light has the longer wavelength, violet or red? Which color of light has the higher energy per photon, violet or red?

9.2 CHLOROPLASTS ■ ■ LEARNING OBJECTIVES 2 3

Diagram the internal structure of a chloroplast and explain how its components interact and facilitate the process of photosynthesis. Describe what happens to an electron in a biological molecule such as chlorophyll when a photon of light energy is absorbed.

If you examine a section of leaf tissue in a microscope, you see that the green pigment, chlorophyll, is not uniformly distributed in

the cell but is confined to organelles called chloroplasts. In plants, chloroplasts lie mainly inside the leaf in the cells of the mesophyll, a layer with many air spaces and a very high concentration of water vapor (FIG. 9-4a). The interior of the leaf exchanges gases with the outside through microscopic pores, called stomata (sing., stoma). Each mesophyll cell has 20 to 100 chloroplasts. The chloroplast, like the mitochondrion, is enclosed by outer and inner membranes (FIG. 9-4b). The inner membrane encloses a fluid-filled region called the stroma, which contains most of the enzymes required to produce carbohydrate molecules. Suspended in the stroma is a third system of membranes that forms an interconnected set of flat, disclike sacs called thylakoids. The thylakoid membrane encloses a fluid-filled interior space, the thylakoid lumen. In some regions of the chloroplast, thylakoid sacs are arranged in stacks called grana (sing., granum). Each granum looks something like a stack of coins, with each “coin” being a thylakoid. Some thylakoid membranes extend from one granum to another. Thylakoid membranes, like the inner mitochondrial membrane (see Chapter 8), are involved in ATP synthesis. (Photosynthetic prokaryotes have no chloroplasts, but thylakoid membranes are often arranged around the periphery of the cell as infoldings of the plasma membrane.)

Chlorophyll is found in the thylakoid membrane Thylakoid membranes contain several kinds of pigments, which are substances that absorb visible light. Different pigments absorb light of different wavelengths. Chlorophyll, the main pigment of photosynthesis, absorbs light primarily in the blue and red regions of the visible spectrum. Green light is not appreciably absorbed by chlorophyll. Plants usually appear green because some of the green light that strikes them is scattered or reflected. A chlorophyll molecule has two main parts, a complex ring and a long side chain (FIG. 9-5). The ring structure, called a porphyrin ring, is made up of joined smaller rings composed of carbon and nitrogen atoms; the porphyrin ring absorbs light energy. The porphyrin ring of chlorophyll is strikingly similar to the heme portion of the red pigment hemoglobin in red blood cells. However, unlike heme, which contains an atom of iron in the center of the ring, chlorophyll contains an atom of magnesium in that position. The chlorophyll molecule also contains a long, hydrocarbon side chain that makes the molecule extremely nonpolar and anchors the chlorophyll in the membrane. All chlorophyll molecules in the thylakoid membrane are associated with specific chlorophyll-binding proteins; biologists have identified about 15 different kinds. Each thylakoid membrane is filled with precisely oriented chlorophyll molecules and chlorophyll-binding proteins that facilitate the transfer of energy from one molecule to another. There are several kinds of chlorophyll. The most important is chlorophyll a, the pigment that initiates the light-dependent reactions of photosynthesis. Chlorophyll b is an accessory pigment that also participates in photosynthesis. It differs from chlorophyll a only in a functional group on the porphyrin ring: the methyl group (OCH3) in chlorophyll a is replaced in chlorophyll b by a

energy passed to it from the light source, or indirectly by energy passed to it from accessory pigments that have become excited by light. When a carotenoid molecule is excited, its energy can be transferred to chlorophyll a. In addition, carotenoids are antioxidants that inactivate highly reactive forms of oxygen generated in the chloroplasts.

terminal carbonyl group (OCHO). This difference shifts the wavelengths of light absorbed and reflected by chlorophyll b, making it appear yellow-green, whereas chlorophyll a appears bright green. Chloroplasts have other accessory photosynthetic pigments, such as carotenoids, which are yellow and orange (see Fig. 3-14). Carotenoids absorb different wavelengths of light than chlorophyll, thereby expanding the spectrum of light that provides energy for photosynthesis. Chlorophyll may be excited by light directly by

Chlorophyll is the main photosynthetic pigment As you have seen, the thylakoid membrane contains more than one kind of pigment. An instrument called a spectrophotometer measures the relative abilities of different pigments to absorb different wavelengths of light. The absorption spectrum of a pigment is a plot of its absorption of light of different wavelengths. FIGURE 9-6a shows the absorption spectra for chlorophylls a and b. in chlorophyll b CH2

CHO

CH

H C

C H 3C

Palisade mesophyll

Porphyrin ring (absorbs light)

C

C

N

N

C

C

N

N

C

C H

Air space

C

Spongy mesophyll

HC

CH2

C

C

O

O

C

CH3

C

C

CH2

CH2CH3

CH

C C

H

C

Mg

HC H 3C

Vein

C

C

C

in chlorophyll a

CH3

C

O

O

O CH3

CH2

Stoma

CH

(a) This leaf cross section reveals that the mesophyll is the photosynthetic tissue. CO2 enters the leaf through tiny pores or stomata, and H2O is carried to the mesophyll in veins.

C

CH3

CH2 CH2

Inner Outer membrane membrane

CH2

Stroma

HC

Hydrocarbon side chain

CH2

E. H. Newcomb and W. P. Wergin, Biological Photo Service

1 μm

CH2 HC

Thylakoid Thylakoid Granum (stack of membrane lumen thylakoids)

(b) In the chloroplast, pigments necessary for the light-capturing reactions of photosynthesis are part of thylakoid membranes, whereas the enzymes for the synthesis of carbohydrate molecules are in the stroma.

FIGURE 9-4 Animated The site of photosynthesis

CH3

CH2 CH2 CH2 CH H 3C

Intermembrane space

CH3

CH2

CH3

FIGURE 9-5 The structure of chlorophyll Chlorophyll consists of a porphyrin ring and a hydrocarbon side chain. The porphyrin ring, with a magnesium atom in its center, contains a system of alternating double and single bonds; these are commonly found in molecules that strongly absorb certain wavelengths of visible light and reflect others (chlorophyll reflects green). Notice that at the top right corner of the diagram, the methyl group (OCH3) distinguishes chlorophyll a from chlorophyll b, which has a carbonyl group (OCHO) in this position. The hydrophobic hydrocarbon side chain anchors chlorophyll to the thylakoid membrane.

An action spectrum of photosynthesis is a graph of the relative effectiveness of different wavelengths of light. To obtain an action spectrum, scientists measure the rate of photosynthesis at each wavelength for leaf cells or tissues exposed to monochromatic light (light of one wavelength) (FIG. 9-6b). In a classic biology experiment, the German biologist T. W. Engelmann obtained the first action spectrum in 1883. Engelmann’s experiment, described in FIGURE 9-7, took advantage of the shape of the chloroplast in a species of the green alga Spirogyra. Its long, filamentous strands are found in freshwater habitats, especially slow-moving or still waters. Spirogyra cells each contain a long, spiral-shaped, emerald-green chloroplast embedded in the cyto-

KEY EXPERIMENT QUESTION: Is a pigment in the chloroplast responsible for photosynthesis? HYPOTHESIS: Engelmann hypothesized that chlorophyll was the main photosynthetic pigment. Accordingly, he predicted that he would observe differences in the amount of photosynthesis, as measured by the amount of oxygen produced, depending on the wavelengths of light used, and that these wavelengths would be consistent with the known absorption spectrum of chlorophyll.

100 80

EXPERIMENT: The photograph (a) shows cells of the filamentous alga Spirogyra, which has a long spiral-shaped chloroplast. The drawing (b) shows how Engelmann used a prism to expose the cells to light that had been separated into various wavelengths. He estimated the formation of oxygen (which he knew was a product of photosynthesis) by exploiting the fact that certain aerobic bacteria would be attracted to the oxygen. As a control (not shown), he also exposed the bacteria to the spectrum of light in the absence of Spirogyra cells.

Chlorophyll b

60 Chlorophyll a

40 20

400

500

600

700

T. E. Adams/Visuals Unlimited

Estimated absorption (%)

plasm. Engelmann exposed these cells to a color spectrum produced by passing light through a prism. He hypothesized that if chlorophyll were indeed responsible for photosynthesis, the process would take place most rapidly in the areas where the chloroplast was illuminated by the colors most strongly absorbed by chlorophyll.

Wavelength (nm)

(a) Chlorophylls a and b absorb light mainly in the blue (422 to 492 nm) and red (647 to 760 nm) regions.

Relative rate of photosynthesis

100

100 µm

(a)

80 60 40 20

380 400

(b)

400

500

600

700

Wavelength (nm)

(b) The action spectrum of photosynthesis indicates the effectiveness of various wavelengths of light in powering photosynthesis. Many plant species have action spectra for photosynthesis that resemble the generalized action spectrum shown here.

500

600

700

RESULTS AND CONCLUSION: Although the bacteria alone (control) showed no preference for any particular wavelength, large numbers were attracted to the photosynthesizing cells in red or blue light, wavelengths that are strongly absorbed by chlorophyll (see Fig. 9-6). Thus, Engelmann concluded that chlorophyll is responsible for photosynthesis.

FIGURE 9-7 Animated The first action spectrum of FIGURE 9-6 A comparison of the absorption spectra for chlorophylls a and b with the action spectrum for photosynthesis

760

Wavelength of light (nm)

photosynthesis

■

■

■

What chloroplast membrane is most important in photosynthesis? What two compartments does it separate? What is the significance of the fact that the combined absorption spectra of chlorophylls a and b roughly match the action spectrum of photosynthesis? Why do they not coincide exactly? Does fluorescence play a role in photosynthesis?

9.3 OVERVIEW OF PHOTOSYNTHESIS ■ ■ LEARNING OBJECTIVES 4 Describe photosynthesis as a redox process. 5 Distinguish between the light-dependent reactions and carbon fixation reactions of photosynthesis.

During photosynthesis, a cell uses light energy captured by chlorophyll to power the synthesis of carbohydrates. The overall reaction of photosynthesis can be summarized as follows: 6 CO 2 + 12 H 2O Carbon dioxide

Water

Light energy

¡ C6 H12O6 + 6 O 2 + 6 H 2O

Chlorophyll

Glucose

Oxygen

6 CO 2 + 6 H2O

Light

¡ C6 H12O6 + 6 O 2

Chlorophyll

When you analyze this process, it appears that hydrogen atoms are transferred from H2O to CO2 to form carbohydrate, so you can recognize it as a redox reaction. Recall from Chapter 7 that in a redox reaction one or more electrons, usually as part of one or more hydrogen atoms, are transferred from an electron donor (a reducing agent) to an electron acceptor (an oxidizing agent). Reduction

6 CO2 + 6 H2O

Light

¡ C 6 H12 O6 + 6 O 2

Chlorophyll

Oxidation

¡

Review

product in others. Furthermore, all the oxygen produced comes from water, so 12 water molecules are required to produce 12 oxygen atoms. However, because there is no net yield of H2O, we can simplify the summary equation of photosynthesis for purposes of discussion:

¡

Yet how could photosynthesis be measured in those technologically unsophisticated days? Engelmann knew that photosynthesis produces oxygen and that certain motile bacteria are attracted to areas of high oxygen concentration. He determined the action spectrum of photosynthesis by observing that the bacteria swam toward the parts of the Spirogyra filaments in the blue and red regions of the spectrum. How did Engelmann know bacteria were not simply attracted to blue or red light? Engelmann exposed bacteria to the spectrum of visible light in the absence of Spirogyra as a control. The bacteria showed no preference for any particular wavelength of light. Because the action spectrum of photosynthesis closely matched the absorption spectrum of chlorophyll, Engelmann concluded that chlorophyll in the chloroplasts (and not another compound in another organelle) is responsible for photosynthesis. Numerous studies using sophisticated instruments have since confirmed Engelmann’s conclusions. If you examine Figure 9-6 closely, you will observe that the action spectrum of photosynthesis does not parallel the absorption spectrum of chlorophyll exactly. This difference occurs because accessory pigments, such as carotenoids, transfer some of the energy of excitation produced by green light to chlorophyll molecules. The presence of these accessory photosynthetic pigments can be demonstrated by chemical analysis of almost any leaf, although it is obvious in temperate climates when leaves change color in the fall. Toward the end of the growing season, chlorophyll breaks down (and its magnesium is stored in the permanent tissues of the tree), leaving orange and yellow accessory pigments in the leaves.

When the electrons are transferred, some of their energy is transferred as well. However, the summary equation of photosynthesis is somewhat misleading because no direct transfer of hydrogen atoms actually occurs. The summary equation describes what happens but not how it happens. The how is more complex and involves multiple steps, many of which are redox reactions. The reactions of photosynthesis are divided into two phases: the light-dependent reactions (the photo part of photosynthesis) and the carbon fixation reactions (the synthesis part of photosynthesis). Each set of reactions occurs in a different part of the chloroplast: the light-dependent reactions in association with the thylakoids, and the carbon fixation reactions in the stroma (FIG. 9-8).

ATP and NADPH are the products of the light-dependent reactions: An overview Light energy is converted to chemical energy in the light-dependent reactions, which are associated with the thylakoids. The lightdependent reactions begin as chlorophyll captures light energy, which causes one of its electrons to move to a higher energy state. The energized electron is transferred to an acceptor molecule and is replaced by an electron from H2O. When this happens, H2O is split and molecular oxygen is released (FIG. 9-9). Some energy of the energized electrons is used to phosphorylate adenosine diphosphate (ADP), forming adenosine triphosphate (ATP). In addition, the coenzyme nicotinamide adenine dinucleotide phosphate (NADP+) becomes reduced, forming NADPH.1 NADPH is a hydrogen carrier similar to NADH, differing by the addition of a phosphate group. Unlike NADH, which is generally associated with catabolic pathways like aerobic cellular respiration, NADPH has the ability to provide high-energy electrons to power

Water

Although the correct way to write the reduced form of NADP+ is NADPH + H+, for simplicity’s sake we present the reduced form as NADPH throughout the book. 1

The equation is typically written in the form just given, with H2O on both sides, because water is a reactant in some reactions and a

KEY POINT

The light-dependent reactions in the thylakoids capture energy as ATP and NADPH, which power the carbon fixation reactions in the stroma. Light-dependent reactions (in thylakoids)

Carbon fixation reactions (in stroma) Chloroplast ATP

Light reactions

ADP NADPH

Calvin cycle

NADP+

H2O

O2

CO2

Carbohydrates

FIGURE 9-8 Animated An overview of photosynthesis

as carbon fixation, these reactions “fix” carbon atoms from CO2 to existing skeletons of organic molecules. Because the carbon fixation reactions have no direct requirement for light, they were previously referred to as the “dark” reactions. However, they do not require darkness; in fact, many of the enzymes involved in carbon fixation are much more active in the light than in the dark. Furthermore, carbon fixation reactions depend on the products of the light-dependent reactions. Carbon fixation reactions take place in the stroma of the chloroplast. Now that we have presented an overview of photosynthesis, let us examine the entire process more closely.

certain reactions in anabolic pathways, such as the carbon fixation reactions of photosynthesis. Thus, the products of the lightdependent reactions, ATP and NADPH, are both needed in the energy-requiring carbon fixation reactions.

Carbohydrates are produced during the carbon fixation reactions: An overview The ATP and NADPH molecules produced during the lightdependent phase are suited for transferring chemical energy but not for long-term energy storage. For this reason, some of their energy is transferred to chemical bonds in carbohydrates, which can be produced in large quantities and stored for future use. Known

Review ■

Bernard Wittich/Visuals Unlimited

■

FIGURE 9-9 Oxygen produced by photosynthesis On sunny days, the oxygen released by aquatic plants is sometimes visible as bubbles in the water. This plant (Elodea) is actively carrying on photosynthesis.

Which is more oxidized, oxygen that is part of a water molecule or molecular oxygen? In what ways do the carbon fixation reactions depend on the lightdependent reactions?

9.4 THE LIGHT-DEPENDENT REACTIONS ■ ■ LEARNING OBJECTIVES 6 Describe the flow of electrons through photosystems I and II in the noncyclic electron transport pathway and the products produced. Contrast this with cyclic electron transport.

7

Explain how a proton (H+) gradient is established across the thylakoid membrane and how this gradient functions in ATP synthesis.

In the light-dependent reactions, the radiant energy from sunlight phosphorylates ADP, producing ATP, and reduces NADP+, forming NADPH. The light energy that chlorophyll captures is

temporarily stored in these two compounds. The light-dependent reactions are summarized as follows: 12 H2O + 12 NADP+ + 18 ADP + 18 Pi

Light

¡ Chlorophyll

6 O2 + 12 NADPH + 18 ATP

Photosystems I and II each consist of a reaction center and multiple antenna complexes The light-dependent reactions of photosynthesis begin when chlorophyll a and/or accessory pigments absorb light. According to the currently accepted model, chlorophylls a and b and accessory pigment molecules are organized with pigment-binding proteins in the thylakoid membrane into units called antenna complexes. The pigments and associated proteins are arranged as highly ordered groups of about 250 chlorophyll molecules associated with specific enzymes and other proteins. Each antenna complex absorbs light energy and transfers it to the reaction center, which consists of chlorophyll molecules and proteins, including electron transfer components, that participate directly in photosynthesis (FIG. 9-10). Light energy is converted to chemical energy in the reaction centers by a series of electron transfer reactions. Two types of photosynthetic units, designated photosystem I and photosystem II, are involved in photosynthesis. Their reaction centers are distinguishable because they are associated with proteins in a way that causes a slight shift in their absorption spectra. Ordinary chlorophyll a has a strong absorption peak at about 660

Primary electron acceptor e-

Chloroplast

Photon

Thylakoid membrane

Photosystem

FIGURE 9-10 Animated Schematic view of a photosystem Chlorophyll molecules (green circles) and accessory pigments (not shown) are arranged in light-harvesting arrays, or antenna complexes. A portion of one such complex within a photosystem is depicted. Each complex consists of several hundred pigment molecules, in association with special proteins (not shown). These proteins hold the pigments in a highly ordered spatial array, such that when a molecule in an antenna complex absorbs a photon, energy derived from that photon is readily passed from one pigment molecule to another (black arrow). When this energy reaches one of the two chlorophyll molecules in the reaction center (green diamonds), an electron becomes energized and is accepted by a primary electron acceptor.

nm. In contrast, the reaction center of photosystem I consists of a pair of chlorophyll a molecules with an absorption peak at 700 nm and is referred to as P700. The reaction center of photosystem II is made up of a pair of chlorophyll a molecules with an absorption peak of about 680 nm and is referred to as P680. When a pigment molecule absorbs light energy, that energy is passed, through a process known as resonance, directly from one pigment molecule to another within the antenna complex until it reaches the reaction center. When the energy reaches a molecule of P700 (in a photosystem I reaction center) or P680 (in a photosystem II reaction center), an electron is then raised to a higher energy level. As we explain in the next section, this energized electron can be donated to an electron acceptor that becomes reduced in the process.

Noncyclic electron transport produces ATP and NADPH Let us begin our discussion of noncyclic electron transport with the events associated with photosystem I (FIG. 9-11). A pigment molecule in an antenna complex associated with photosystem I absorbs a photon of light. The absorbed energy is transferred from one pigment molecule to another until it reaches the reaction center, where it excites an electron in a molecule of P700. This energized electron is transferred to a primary electron acceptor, a special molecule of chlorophyll a, which is the first of several electron acceptors in a series. The energized electron is passed along an electron transport chain from one electron acceptor to another, until it is passed to ferredoxin, an iron-containing protein. Ferredoxin transfers the electron to NADP+ in the presence of the enzyme ferredoxin–NADP+ reductase. When NADP+ accepts two electrons, they unite with a proton + (H ); thus, the reduced form of NADP+ is NADPH, which is released into the stroma. P700 becomes positively charged when it gives up an electron to the primary electron acceptor; the missing electron is replaced by one donated by photosystem II. As in photosystem I, photosystem II becomes activated when a pigment molecule in an antenna complex absorbs a photon of light energy. The energy is transferred to the reaction center, where it causes an electron in a molecule of P680 to move to a higher energy level. This energized electron is accepted by a primary electron acceptor (a highly modified chlorophyll molecule known as pheophytin) and then passes along an electron transport chain until it is donated to P700 in photosystem I. How is the electron that has been donated to the electron transport chain replaced? This occurs through photolysis (light splitting) of water, a process that not only yields electrons but also is the source of almost all the oxygen in Earth’s atmosphere. A molecule of P680 that has given up an energized electron to the primary electron acceptor is positively charged (P680+). P680+ is an oxidizing agent so strong that it pulls electrons away from an oxygen atom that is part of an H2O molecule. In a reaction catalyzed by a unique, manganese-containing enzyme, water is broken into its components: two electrons, two protons, and oxygen. Each electron is donated to a P680 molecule, which then loses its positive charge; the protons are released into the thylakoid lumen. Because oxygen does not exist in atomic form, the oxygen produced

KEY POINT

Noncyclic electron transport converts light energy to chemical energy in ATP and NADPH.

Light-dependent reactions (in thylakoids)

Carbon fixation reactions (in stroma)

Chloroplast ATP Light reactions

ADP

Calvin cycle

NADPH NADP

H2O

2e–

2e–

Primary electron acceptor

Primary electron acceptor

Electron transport chain Production of ATP by chemiosmosis

Relative energy level

Carbohydrates

CO2

O2

Electron transport chain

Ferredoxin 2e–

H+ (from medium)

ATP

NADP+

NADPH

2e– +

1/2 O2 + 2 H

2e–

Photosystem I (P700)

H2O Photosystem II (P680)

1

Electrons are supplied to system from the splitting of H2O by photosystem II, with release of O2 as byproduct. When photosystem II is activated by absorbing photons, electrons are passed along the electron transport chain and are eventually donated to photosystem I.

2

Electrons in photosystem I are “re-energized“ by absorption of additional light energy and are passed to NADP+, forming NADPH.

FIGURE 9-11 Animated Noncyclic electron transport In noncyclic electron transport, the formation of ATP is coupled to one-way flow of energized electrons (orange arrows) from H2O (lower left) to NADP+ (far right). Single electrons actually pass down the electron transport chain; two are shown in this figure because two electrons are required to form one molecule of NADPH.

by splitting one H2O molecule is written –12 O2. Two water molecules must be split to yield one oxygen molecule. The photolysis of water is a remarkable reaction, but its name is somewhat misleading because it implies that water is broken by light. Actually, light splits water indirectly by causing P680 to become oxidized. Noncyclic electron transport is a continuous linear process In the presence of light, there is a continuous, one-way flow of electrons from the ultimate electron source, H2O, to the terminal electron acceptor, NADP+. Water undergoes enzymatically catalyzed photolysis to replace energized electrons donated to the electron

transport chain by molecules of P680 in photosystem II. These electrons travel down the electron transport chain that connects photosystem II with photosystem I. Thus, they provide a continuous supply of replacements for energized electrons that have been given up by P700. As electrons are transferred along the electron transport chain that connects photosystem II with photosystem I, they lose energy. Some of the energy released is used to pump protons across the thylakoid membrane, from the stroma to the thylakoid lumen, producing a proton gradient. The energy of this proton gradient is harnessed to produce ATP from ADP by chemiosmosis, which we discuss later in the chapter. ATP and NADPH, the products of

the light-dependent reactions, are released into the stroma, where both are required by the carbon fixation reactions.

Cyclic electron transport produces ATP but no NADPH Only photosystem I is involved in cyclic electron transport, the simplest light-dependent reaction. The pathway is cyclic because energized electrons that originate from P700 at the reaction center eventually return to P700. In the presence of light, electrons flow continuously through an electron transport chain within the thylakoid membrane. As they pass from one acceptor to another, the electrons lose energy, some of which is used to pump protons across the thylakoid membrane. An enzyme (ATP synthase) in the thylakoid membrane uses the energy of the proton gradient to manufacture ATP. NADPH is not produced, H2O is not split, and oxygen is not generated. By itself, cyclic electron transport could not serve as the basis of photosynthesis because, as we explain later in the chapter, NADPH is required to reduce CO2 to carbohydrate. The significance of cyclic electron transport to photosynthesis in plants is unclear. Cyclic electron transport may occur in plant cells when there is too little NADP+ to accept electrons from ferredoxin. There is evidence that cyclic electron flow may help maintain the optimal ratio of ATP to NADPH required for carbon fixation, as well as provide extra ATP to power other ATP-requiring processes in chloroplasts. Biologists generally agree that ancient bacteria used this process to produce ATP from light energy. A reaction pathway analogous to cyclic electron transport in plants is present in some modern photosynthetic prokaryotes. Noncyclic and cyclic electron transport are compared in TABLE 9-1.

reactions, losing some of its energy at each step. Some of the energy given up by the electron is not lost by the system, however; it is used to provide energy for ATP synthesis. Because the synthesis of ATP (that is, the phosphorylation of ADP) is coupled to the transport of electrons that have been energized by photons of light, the process is called photophosphorylation. The chemiosmotic model explains the coupling of ATP synthesis and electron transport As discussed earlier, the pigments and electron acceptors of the light-dependent reactions are embedded in the thylakoid membrane. Energy released from electrons traveling through the chain of acceptors is used to pump protons from the stroma, across the thylakoid membrane, and into the thylakoid lumen (FIG. 9-12). Thus, the pumping of protons results in the formation of a proton gradient across the thylakoid membrane. Protons also accumulate in the thylakoid lumen as water is split during noncyclic electron transport. Because protons are actually hydrogen ions (H+), the accumulation of protons causes the pH of the thylakoid interior to fall to a pH of about 5 in the thylakoid lumen, compared to a pH of about 8 in the stroma. This difference of about 3 pH units across the thylakoid membrane means there is an approximately thousand-fold difference in hydrogen ion concentration. The proton gradient has a great deal of free energy because of its state of low entropy. How does the chloroplast convert that energy to a more useful form? According to the general principles of diffusion, the concentrated protons inside the thylakoid might be expected to diffuse out readily. However, they are prevented from doing so because the thylakoid membrane is impermeable to H+

ATP synthesis occurs by chemiosmosis Stroma

Each member of the electron transport chain that links photosystem II to photosystem I can exist in an oxidized (lower-energy) form and a reduced (higher-energy) form. The electron accepted from P680 by the primary electron acceptor is highly energized; it is passed from one carrier to the next in a series of exergonic redox

TABLE 9-1

Thylakoid lumen

A Comparison of Noncyclic and Cyclic Electron Transport Noncyclic Electron Transport

Cyclic Electron Transport

Electron source

H 2O

Oxygen released? Terminal electron acceptor Form in which energy is temporarily captured Photosystem(s) required

Yes (from H2O) NADP+

None—electrons cycle through the system No None—electrons cycle through the system ATP (by chemiosmosis)

ATP (by chemiosmosis); NADPH PS I (P700) and PS II (P680)

Thylakoid membrane

Protons (H+)

FIGURE 9-12 The accumulation of protons in the thylakoid lumen PS I (P700) only

As electrons move down the electron transport chain, protons (H+) move from the stroma to the thylakoid lumen, creating a proton gradient. The greater concentration of H+ in the thylakoid lumen lowers the pH.

except through certain channels formed by the enzyme ATP synthase. This enzyme, a transmembrane protein also found in mitochondria, forms complexes so large they can be seen in electron micrographs (see Fig. 8-11b). ATP synthase complexes project into the stroma. As the protons diffuse through an ATP synthase complex, free energy decreases as a consequence of an increase in entropy. Each ATP synthase complex couples this exergonic process of diffusion down a concentration gradient to the endergonic process of phosphorylation of ADP to form ATP, which is released

KEY POINT

into the stroma (FIG. 9-13). The movement of protons through ATP synthase is thought to induce changes in the conformation of the enzyme that are necessary for the synthesis of ATP. It is estimated that for every four protons that move through ATP synthase, one ATP molecule is synthesized. The mechanism by which the phosphorylation of ADP is coupled to diffusion down a proton gradient is called chemiosmosis. As the essential connection between the electron transport chain and the phosphorylation of ADP, chemiosmosis is a basic

Electron carriers associated with the thylakoid membrane transfer energized electrons from water to NADP+, forming NADPH. ATP is generated by chemiosmosis.

Light-dependent reactions

Carbon fixation reactions

Chloroplast ATP ADP

Light reactions

NADPH

Calvin cycle

NADP

H2O

CO2

O2

Carbohydrates

Thylakoid membrane

Thylakoid lumen

H+

H+

H+

H

1/2 O O22 + 2 H+

H2O

H

H+

H

H

Plastocyanin

H H

+

H

+

Plastoquinone

+

+

H

+

+

H

H

+

H

+

H+

+

Photon

H+

H+

+

+

H

H+ H+

+

3

H+

ATP synthase

Photon Thylakoid Photosystem II membrane

Photosystem I

FerredoxinNADP+ reductase

1

H+

Cytochrome complex 2

Ferredoxin H+

4

+ NADP+

NADPH

ADP + Pi ATP

Stroma

1

Orange arrows indicate pathway of electrons along electron transport chain in thylakoid membrane. Electron carriers within membrane become alternately reduced and oxidized as they accept and donate electrons.

2

Energy released during electron transport is used to transport H+ from the stroma to the thylakoid lumen, where a high concentration of H+ accumulates.

FIGURE 9-13 A detailed look at electron transport and chemiosmosis

3

H+ are prevented from diffusing back into stroma except through special channels in ATP synthase in the thylakoid membrane.

4

H+ flows through ATP synthase, generating ATP.

TABLE 9-2

A Comparison of Photosynthesis and Aerobic Respiration Photosynthesis

Aerobic Respiration

Sites involved (in eukaryotic cells) ATP production

Anabolism CO2, H2O C6H12O6, O2 Cells that contain chlorophyll (certain cells of plants, algae, and some bacteria) Chloroplasts By photophosphorylation (a chemiosmotic process)

Principal electron transfer compound Location of electron transport chain Source of electrons for electron transport chain Terminal electron acceptor for electron transport chain

NADP+ is reduced to form NADPH* Thylakoid membrane In noncyclic electron transport: H2O (undergoes photolysis to yield electrons, protons, and oxygen) In noncyclic electron transport: NADP+ (becomes reduced to form NADPH)

Catabolism C6H12O6, O2 CO2, H2O Every actively metabolizing cell has aerobic respiration or some other energy-releasing pathway Cytosol (glycolysis); mitochondria By substrate-level phosphorylation and by oxidative phosphorylation (a chemiosmotic process) NAD+ is reduced to form NADH* Mitochondrial inner membrane (cristae) Immediate source: NADH, FADH2 Ultimate source: glucose or other carbohydrate O2 (becomes reduced to form H2O)

Type of metabolic reaction Raw materials End products Which cells have these processes?

*NADPH and NADH are very similar hydrogen (i.e., electron) carriers, differing only in a single phosphate group. However, NADPH generally works with enzymes in anabolic pathways, such as photosynthesis. NADH is associated with catabolic pathways, such as cellular respiration.

mechanism of energy coupling in cells. You may recall from Chapter 8 that chemiosmosis also occurs in aerobic respiration (see TABLE 9-2).

Review ■ ■ ■

Why is molecular oxygen a necessary byproduct of photosynthesis? What process is the actual mechanism of photophosphorylation? Why are both photosystems I and II required for photosynthesis? Can cyclic phosphorylation alone support photosynthesis? Explain your answer.

9.5 THE CARBON FIXATION REACTIONS ■ ■ LEARNING OBJECTIVES 8 Summarize the three phases of the Calvin cycle and indicate the roles of ATP and NADPH in the process.

9 Discuss how photorespiration reduces photosynthetic efficiency. 10 Compare the C4 and CAM pathways.

In carbon fixation, the energy of ATP and NADPH is used in the formation of organic molecules from CO2. The carbon fixation reactions may be summarized as follows: 12 NADPH + 18 ATP + 6 CO2 ¡ C6H12O6 + 12 NADP+ + 18 ADP + 18 Pi + 6 H2O

Most plants use the Calvin cycle to fix carbon Carbon fixation occurs in the stroma through a sequence of 13 reactions known as the Calvin cycle. During the 1950s, University of California researchers Melvin Calvin, Andrew Benson, and oth-

ers elucidated the details of this cycle. Calvin was awarded a Nobel Prize in Chemistry in 1961. The 13 reactions of the Calvin cycle are divided into three phases: CO2 uptake, carbon reduction, and RuBP regeneration (FIG. 9-14). All 13 enzymes that catalyze steps in the Calvin cycle are located in the stroma of the chloroplast. Ten of the enzymes also participate in glycolysis (see Chapter 8). These enzymes catalyze reversible reactions, degrading carbohydrate molecules in cellular respiration and synthesizing carbohydrate molecules in photosynthesis. 1. CO2 uptake. The first phase of the Calvin cycle consists of a single reaction in which a molecule of CO2 reacts with a phosphorylated five-carbon compound, ribulose bisphosphate (RuBP). This reaction is catalyzed by the enzyme ribulose bisphosphate carboxylase/oxygenase, also known as rubisco. The chloroplast contains more rubisco enzyme than any other protein, and rubisco may be one of the most abundant proteins in the biosphere. The product of this reaction is an unstable six-carbon intermediate, which immediately breaks down into two molecules of phosphoglycerate (PGA) with three carbons each. The carbon that was originally part of a CO2 molecule is now part of a carbon skeleton; the carbon has been “fixed.” The Calvin cycle is also known as the C3 pathway because the product of the initial carbon fixation reaction is a three-carbon compound. Plants that initially fix carbon in this way are called C3 plants. 2. Carbon reduction. The second phase of the Calvin cycle consists of two steps in which the energy and reducing power from ATP and NADPH (both produced in the light-dependent reactions) are used to convert the PGA molecules to glyceraldehyde-3-phosphate (G3P). As shown in Figure 9-14, for every six carbons that enter the cycle as CO2, six carbons can leave the system as two molecules of G3P, to be used in car-

KEY POINT

ATP and NADPH provide the energy that drives carbon fixation in the Calvin cycle.

Light-dependent reactions

Carbon fixation reactions

Chloroplast ATP Light reactions

ADP NADPH

Calvin cycle

6 molecules of CO2

NADP

H2O

O2

CO2

CO2 molecules are captured by RuBP, resulting in unstable intermediate that is immediately broken apart into 2 PGA.

Carbohydrates

6 molecules of ribulose bisphosphate (RuBP) P

P

12 molecules of phosphoglycerate (PGA)

1

CO2 uptake phase

6 ADP

P 6

ATP

12 3

6 molecules of ribulose phosphate (RP) P

RuBP regeneration phase

10 molecules of G3P

CALVIN CYCLE

ATP

12 ADP 2

Carbon reduction phase

P

12 NADPH

12 NADP++

Glucose and other carbohydrate synthesis

12 Pi P 12 molecules of glyceraldehyde-3phosphate (G3P) P 2 molecules of glyceraldehyde-3phosphate (G3P)

FIGURE 9-14 Animated A detailed look at the Calvin cycle 1 This diagram, in which carbon atoms are black balls, shows that six ●

molecules of CO2 must be “fixed” (incorporated into pre-existing carbon skeletons) in the CO2 uptake phase to produce one molecule of a sixcarbon sugar such as glucose. ● 2 Glyceraldehyde-3-phosphate (G3P) is formed in the carbon reduction phase. Two G3P molecules “leave”

bohydrate synthesis. Each of these three-carbon molecules of G3P is essentially half a hexose (six-carbon sugar) molecule. (In fact, you may recall that G3P is a key intermediate in the splitting of sugar in glycolysis; see Figs. 8-3 and 8-4.)

Through a series of reactions G3P is rearranged into new RuBP molecules or another sugar.

PGA is phosphorylated by ATP and reduced by NADPH. Removal of phosphate results in formation of G3P.

the cycle for every glucose formed. ● 3 Ribulose bisphosphate (RuBP) is regenerated, and a new cycle can begin. Although these reactions do not require light directly, the energy that drives the Calvin cycle comes from ATP and NADPH, which are the products of the light-dependent reactions.

The reaction of two molecules of G3P is exergonic and leads to the formation of glucose or fructose. In some plants, glucose and fructose are then joined to produce sucrose (common table sugar). (Sucrose can be harvested from sugarcane,

TABLE 9-3

Summary of Photosynthesis

Reaction Series

Summary of Process

Light-dependent reactions (take place in thylakoid membranes) Photochemical reactions

Energy from sunlight used to split water, manufacture ATP, and reduce NADP+

Electron transport

Chemiosmosis

Carbon fixation reactions (take place in stroma)

Needed Materials

End Products

Chlorophyll-activated; reaction center gives up photoexcited electron to electron acceptor Electrons transported along chain of electron acceptors in thylakoid membranes; electrons reduce NADP+; splitting of water provides some H+ that accumulates inside thylakoid space H+ permitted to diffuse across the thylakoid membrane down their gradient; they cross the membrane through special channels in ATP synthase complex; energy released is used to produce ATP

Light energy; pigments (chlorophyll) Electrons, NADP+, H2O, electron acceptors

Electrons

Proton gradient, ADP + Pi, ATP synthase

ATP

Carbon fixation: carbon dioxide used to make carbohydrate

Ribulose bisphosphate, CO2, ATP, NADPH, necessary enzymes

Carbohydrates, ADP + Pi, NADP+

sugar beets, and maple sap.) The plant cell also uses glucose to produce starch or cellulose. 3. RuBP regeneration. Notice that although 2 G3P molecules are removed from the cycle, 10 G3P molecules remain; this represents a total of 30 carbon atoms. Through a series of 10 reactions that make up the third phase of the Calvin cycle, these 30 carbons and their associated atoms become rearranged into six molecules of ribulose phosphate, each of which becomes phosphorylated by ATP to produce RuBP, the five-carbon compound with which the cycle started. These RuBP molecules begin the process of CO2 fixation and eventual G3P production once again. In summary, the inputs required for the carbon fixation reactions are six molecules of CO2 (the source of both the carbons and the oxygens in carbohydrate), phosphates transferred from ATP, and electrons (as hydrogen) provided by NADPH (but ultimately derived from the photolysis of water). In the end, the six carbons from the CO2 are accounted for by the harvest of a hexose molecule. The remaining G3P molecules are used to synthesize the RuBP molecules with which more CO2 molecules may combine. TABLE 9-3 provides a summary of photosynthesis.

Photorespiration reduces photosynthetic efficiency Many C3 plants, including certain agriculturally important crops such as soybeans, wheat, and potatoes, do not yield as much carbohydrate from photosynthesis as might be expected, especially during periods of very hot temperature in summer. This phenomenon is a consequence of trade-offs between the plant’s need for CO2 and its need to prevent water loss. Recall that most photosynthesis occurs in mesophyll cells inside the leaf and that the entry and exit of gases from the interior of the leaf are regulated by stomata, tiny pores concentrated on the underside of the leaf (see Fig. 9-4a). On hot, dry days, plants close their stomata to conserve

NADPH, O2

water. Once the stomata close, photosynthesis rapidly uses up the CO2 remaining in the leaf and produces O2, which accumulates in the chloroplasts. Recall that the enzyme RuBP carboxylase/oxygenase (rubisco) catalyzes CO2 fixation in the Calvin cycle by attaching CO2 to RuBP. As its full name implies, rubisco acts not only as a carboxylase but also as an oxygenase because high levels of O2 compete with CO2 for the active site of rubisco. Some of the intermediates involved in the Calvin cycle are degraded to CO2 and H2O in a process that is called photorespiration because (1) it occurs in the presence of light; and as in aerobic respiration, (2) it requires oxygen and (3) produces CO2 and H2O. However, photorespiration does not produce ATP, and it reduces photosynthetic efficiency because it removes some of the intermediates used in the Calvin cycle. The reasons for photorespiration are incompletely understood, although scientists hypothesize that it reflects the origin of rubisco at an ancient time when CO2 levels were high and molecular oxygen levels were low. This view is supported by recent evidence that some amino acid sequences in rubisco are similar to sequences in certain bacterial proteins that apparently evolved prior to the evolution of the Calvin cycle. Genetic engineering to produce plants with rubisco that has a much lower affinity for oxygen is a promising area of research to improve yields of certain valuable crop plants.

The initial carbon fixation step differs in C4 plants and in CAM plants Photorespiration is not the only problem faced by plants engaged in photosynthesis. Because CO2 is not a very abundant gas (composing only 0.038% of the atmosphere), it is not easy for plants to obtain the CO2 they need. As you have learned, when conditions are hot and dry, the stomata close to reduce the loss of water vapor, greatly diminishing the supply of CO2. Ironically, CO2 is potentially less available at the very times when maximum sunlight is available to power the light-dependent reactions.

Many plant species living in hot, dry environments have adaptations that facilitate carbon fixation. C4 plants first fix CO2 into a four-carbon compound, oxaloacetate. CAM plants initially fix carbon at night through the formation of oxaloacetate. These special pathways found in C4 and CAM plants precede the Calvin cycle (C3 pathway); they do not replace it. The C4 pathway efficiently fixes CO2 at low concentrations The C4 pathway, in which CO2 is fixed through the formation of oxaloacetate, occurs not only before the C3 pathway but also in different cells. Leaf anatomy is usually distinctive in C4 plants. The photosynthetic mesophyll cells are closely associated with prominent, chloroplast-containing bundle sheath cells, which tightly encircle the veins of the leaf (FIG. 9-15). The C4 pathway occurs in the mesophyll cells, whereas the Calvin cycle takes place within the bundle sheath cells. The key component of the C4 pathway is a remarkable enzyme that has an extremely high affinity for CO2, binding it effectively even at unusually low concentrations. This enzyme, PEP carboxylase, catalyzes the reaction by which CO2 reacts with the three-carbon compound phosphoenolpyruvate (PEP), forming oxaloacetate (FIG. 9-16). In a step that requires NADPH, oxaloacetate is converted to some other four-carbon compound, usually malate. The malate then passes to chloroplasts within bundle sheath cells, where a different enzyme catalyzes the decarboxylation of malate to yield pyruvate (which has three carbons) and CO2. NADPH is formed, replacing the one used earlier. malate + NADP+ ¡ pyruvate + CO2 + NADPH

The CO2 released in the bundle sheath cell combines with ribulose bisphosphate in a reaction catalyzed by rubisco and goes through the Calvin cycle in the usual manner. The pyruvate formed in the decarboxylation reaction returns to the mesophyll cell, where it reacts with ATP to regenerate phosphoenolpyruvate. Because the C4 pathway captures CO2 and provides it to the bundle sheath cells so efficiently, CO2 concentration within the bundle sheath cells is about 10 to 60 times as great as its concentration in the mesophyll cells of plants having only the C3 pathway. Photorespiration is negligible in C4 plants such as crabgrass because the concentration of CO2 in bundle sheath cells (where rubisco is present) is always high. The combined C3–C4 pathway involves the expenditure of 30 ATPs per hexose, rather than the 18 ATPs used by the C3 pathway alone. The extra energy expense required to regenerate PEP from pyruvate is worthwhile at high light intensities because it ensures a high concentration of CO2 in the bundle sheath cells and permits them to carry on photosynthesis at a rapid rate. At lower light intensities and temperatures, C3 plants are favored. For example, winter rye, a C3 plant, grows lavishly in cool weather, when crabgrass cannot because it requires more energy to fix CO2.

CAM plants fix CO2 at night Plants living in dry, or xeric, conditions have a number of structural adaptations that enable them to survive. Many xeric plants have physiological adaptations as well, including a special carbon fixation pathway, the crassulacean acid metabolism (CAM) pathway. The name comes from the stonecrop plant family (the Crassulaceae), which uses the CAM pathway, although the pathway has evolved

Upper epidermis Palisade mesophyll Bundle sheath cells of veins Mesophyll

Spongy mesophyll Chloroplasts

(a) In C3 plants, the Calvin cycle takes place in the mesophyll cells and the bundle sheath cells are nonphotosynthetic.

FIGURE 9-15 C3 and C4 plant structure compared

independently in some members of more than 25 other plant families, including the cactus family (Cactaceae), the lily family (Liliaceae), and the orchid family (Orchidaceae) (FIG. 9-17). Unlike most plants, CAM plants open their stomata at night, admitting CO2 while minimizing water loss. They use the enzyme PEP carboxylase to fix CO2, forming oxaloacetate, which is converted to malate and stored in cell vacuoles. During the day, when stomata are closed and gas exchange cannot occur between the plant and the atmosphere, CO2 is removed from malate by a decarboxylation reaction. Now the CO2 is available within the leaf tissue to be fixed into sugar by the Calvin cycle (C3 pathway). The CAM pathway is very similar to the C4 pathway but with important differences. C4 plants initially fix CO2 into four-carbon organic acids in mesophyll cells. The acids are later decarboxylated to produce CO2, which is fixed by the C3 pathway in the bundle sheath cells. In other words, the C4 and C3 pathways occur in different locations within the leaf of a C4 plant. In CAM plants, the initial fixation of CO2 occurs at night. Decarboxylation of malate and subsequent production of sugar from CO2 by the normal C3 photosynthetic pathway occur during the day. In other words, the CAM and C3 pathways occur at different times within the same cell of a CAM plant. Although it does not promote rapid growth the way that the C4 pathway does, the CAM pathway is a very successful adaptation to xeric conditions. CAM plants can exchange gases for photosynthesis and reduce water loss significantly. Plants with CAM photosynthesis survive in deserts where neither C3 nor C4 plants can.

CO2 Mesophyll cell

(3C)

Phosphoenolpyruvate

Oxaloacetate (4C) NADPH

ADP

+

NADP Malate (4C)

ATP Pyruvate (3C)

(3C) Pyruvate

Malate (4C)

Glucose

C

NADP+ CO2 alvin

Bundle sheath cell

NADPH

c y c le

Vein

FIGURE 9-16 Animated Summary of the C4 pathway CO2 combines with phosphoenolpyruvate (PEP) in the chloroplasts of mesophyll cells, forming a four-carbon compound that is converted to malate. Malate goes to the chloroplasts of bundle sheath cells, where it is decarboxylated. The CO2 released in the bundle sheath cell is used to make carbohydrate by way of the Calvin cycle.

Review ■ ■ ■ ■

What are the three phases of the Calvin cycle? Which phase of the Calvin cycle requires both ATP and NADPH? In what ways does photorespiration differ from aerobic respiration? Do C3, C4, and CAM plants all have rubisco? PEP carboxylase?

9.6 METABOLIC DIVERSITY ■ ■ LEARNING OBJECTIVE Robert W. Domm/Visuals Unlimited

11 Contrast photoautotrophs and chemoheterotrophs with respect to their

FIGURE 9-17 A typical CAM plant Prickly pear cactus (Opuntia) is a CAM plant. The more than 200 species of Opuntia living today originated in various xeric habitats in North and South America.

energy and carbon sources.

Land plants, algae, and certain prokaryotes are known as photoautotrophs. They are phototrophs because they use light energy to make ATP and NADPH, which temporarily hold chemical energy but are unstable and cannot be stockpiled in the cell. They are autotrophs (from the Greek auto, which means “self,” and trophos, which means “nourishing”) that synthesize complex organic compounds from simpler, inorganic raw materials. The chemical energy of ATP and NADPH then drives carbon fixation, the anabolic pathway in which stable organic molecules are synthesized from CO2 and water. These organic compounds are used not only as starting

materials to synthesize all the other organic compounds the photosynthetic organism needs (such as complex carbohydrates, amino acids, and lipids) but also for energy storage. Glucose and other carbohydrates produced during photosynthesis are relatively reduced compounds that can be subsequently oxidized by aerobic respiration or by some other catabolic pathway (see Chapter 8). In contrast, animals, fungi, and most bacteria are known as chemoheterotrophs. They are chemotrophs because they obtain energy from chemicals, typically by redox reactions (see Chapters 7 and 8). They are heterotrophs (from the Greek heter, which means “other,” and trophos, which means “nourishing”) because they cannot fix carbon; they use organic molecules produced by other organisms as the building blocks from which they synthesize the carbon compounds they need. We are so familiar with plants as photoautotrophs and animals such as ourselves as chemoheterotrophs that we tend to think that all organisms should fit in these two “mainstream” categories. Two other types of nutrition are found in certain bacteria. A few bacteria, known as nonsulfur purple bacteria, are photoheterotrophs, able to use light energy but unable to carry out carbon fixation, so they must obtain carbon from organic compounds. Some other bacteria are chemoautotrophs, which obtain their energy from the oxidation of reduced inorganic molecules such as hydrogen sulfide (H2S), nitrite (NO2−), or ammonia (NH3). Some of this captured energy is subsequently used to carry out carbon fixation.

Review ■

How does a green plant obtain energy? carbon? How does your body obtain these things?

9.7 PHOTOSYNTHESIS IN PLANTS AND IN THE ENVIRONMENT ■ ■ LEARNING OBJECTIVE 12 State the importance of photosynthesis both in a plant and to other organisms.

Although we characterize plants as photoautotrophs, not all plant cells carry out photosynthesis, and even cells with chloroplasts also possess mitochondria and carry out aerobic respiration. In fact, respiration utilizing the organic molecules the plant has made for itself is the direct source of ATP needed for most plant metabolism. Several mechanisms regulate the relative activities of photosynthesis and aerobic respiration in plants. Although the enzymes of the Calvin cycle do not require light to function, they are actually regulated by light. As a consequence of the light-requiring reactions, the stroma becomes more basic (approximately pH 8), activating rubisco and other Calvin cycle enzymes. In contrast, light tends to inhibit the enzymes of glycolysis in the cytosol. Hence photosynthesis, not aerobic respiration, is favored in the light. When light is very dim, at a point known as the light com-

pensation point, photosynthesis still occurs, but it is not evident because the rate of CO2 fixation by photosynthesis is equal to the rate of CO2 release through aerobic respiration. On the other hand, when light is very bright, photorespiration can significantly diminish photosynthetic yields. As we have seen in this chapter, the reactions of the Calvin cycle provide a net yield of the three-carbon phosphorylated sugar, G3P. What are the various fates of G3P in the plant? Consider a leaf cell actively conducting photosynthesis. A series of enzymes may convert some of the G3P to glucose and then to starch. This starch is stored in starch granules that form inside chloroplasts. It has been recently shown that when this starch is broken down, the disaccharide maltose is typically formed (see Fig. 3-8a). Maltose is transported out of the chloroplast and then cleaved in the cytosol, providing glucose for aerobic respiration. Not all G3P ends up as carbohydrate; some is ultimately converted to amino acids, fatty acids, and other organic molecules needed by the photosynthetic cell. Some of the G3P is exported to the cytosol, where enzymes convert it to the disaccharide sucrose (see Fig. 3-8b). Sucrose is then actively transported out of the cell, moves through the vascular system of the plant (see Chapter 35 for a discussion of plant transport), and is actively transported into the various cells. Sucrose can be broken down to glucose and fructose, which are used in aerobic respiration or as starting points for the synthesis of the various organic molecules the cells need, such as amino acids, lipids, and carbohydrates. Important carbohydrates include cellulose for cell walls (see Fig. 3-10) and starch, particularly in starch-storing structures such as roots (see Fig. 3-9a) and developing seeds and tubers (such as potatoes). The benefits of photosynthesis in the environment are staggering. Of course, by fixing carbon, photoautotrophs are the ultimate source of virtually all organic molecules used as energy and carbon sources by chemoheterotrophs such as ourselves (for an exception, see Inquiring About: Life without the Sun in Chapter 55). In carrying out carbon fixation, photoautotrophs remove CO2 from the atmosphere, thereby slowing global warming (see Chapter 57). Also of prime importance is the fact that photolysis of water by photosystem II releases the O2 that all aerobic organisms require for aerobic respiration. Molecular oxygen is so reactive that it could not be maintained in the atmosphere if it were not constantly replenished in this way. As discussed in Chapter 21, the evolution of oxygen-producing photosynthesis was a critical event in the history of life on Earth, which not only permitted the evolution of aerobic organisms, but also made terrestrial life possible because in the stratosphere O2 is converted to ozone (O3), which shields the planet from damaging ultraviolet light.

Review ■ ■

How does a root cell obtain energy? organic molecules? What is the source of molecular oxygen in Earth’s atmosphere?

■ ■

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

9.1 (page 194) 1 Describe the physical properties of light and explain the relationship between a wavelength of light and its energy. ■ Light consists of particles called photons that move as waves. ■ Photons with shorter wavelengths have more energy than those with longer wavelengths.

■

■

9.2 (page 195) 2 Diagram the internal structure of a chloroplast and explain how its components interact and facilitate the process of photosynthesis. In plants, photosynthesis occurs in chloroplasts, which are located mainly within mesophyll cells inside the leaf. ■ Chloroplasts are organelles enclosed by a double membrane; the inner membrane encloses the stroma in which membranous, saclike thylakoids are suspended. Thylakoids enclose the thylakoid lumen. Thylakoids arranged in stacks are called grana. ■ Chlorophyll a, chlorophyll b, carotenoids, and other photosynthetic pigments are components of the thylakoid membranes of chloroplasts. 3 Describe what happens to an electron in a biological molecule such as chlorophyll when a photon of light energy is absorbed. ■ Photons excite biological molecules such as chlorophyll and other photosynthetic pigments, causing one or more electrons to become energized. These energized electrons may be accepted by electron acceptor compounds. ■ The combined absorption spectra of chlorophylls a and b are similar to the action spectrum for photosynthesis.

■

■

9.3 (page 198) 4 Describe photosynthesis as a redox process. During photosynthesis, light energy is captured and converted to the chemical energy of carbohydrates; hydrogens from water are used to reduce carbon, and oxygen derived from water becomes oxidized, forming molecular oxygen. 5 Distinguish between the light-dependent reactions and carbon fixation reactions of photosynthesis. ■ In the light-dependent reactions, electrons energized by light are used to generate ATP and NADPH; these compounds provide energy for the formation of carbohydrates during the carbon fixation reactions. ■

Light-dependent reactions (in thylakoids)

Carbon fixation reactions (in stroma) Chloroplast ATP

Light reactions

ADP NADPH

Calvin cycle

NADP+

H2O

O2

CO2

Carbohydrates

■

■

Experience the process of noncyclic electron transport by clicking on the figure in CengageNOW.

7 Explain how a proton (H+) gradient is established across the thylakoid membrane and how this gradient functions in ATP synthesis. ■ Photophosphorylation is the synthesis of ATP coupled to the transport of electrons energized by photons of light. Some of the energy of the electrons is used to pump protons across the thylakoid membrane, providing the energy to generate ATP by chemiosmosis. ■ As protons diffuse through ATP synthase, an enzyme complex in the thylakoid membrane, ADP is phosphorylated to form ATP.

9.5 (page 204) 8 Summarize the three phases of the Calvin cycle and indicate the roles of ATP and NADPH in the process. ■ The carbon fixation reactions proceed by way of the Calvin cycle, also known as the C3 pathway. ■ In the CO2 uptake phase of the Calvin cycle, CO2 is combined with ribulose bisphosphate (RuBP), a five-carbon sugar, by the enzyme ribulose bisphosphate carboxylase/oxygenase, commonly known as rubisco, forming the three-carbon molecule phosphoglycerate (PGA). In the carbon reduction phase of the Calvin cycle, the energy and ■ reducing power of ATP and NADPH are used to convert PGA molecules to glyceraldehyde-3-phosphate (G3P). For every 6 CO2 molecules fixed, 12 molecules of G3P are produced, and 2 molecules of G3P leave the cycle to produce the equivalent of 1 molecule of glucose. ■ In the RuBP regeneration phase of the Calvin cycle, the remaining G3P molecules are modified to regenerate RuBP. See the Calvin cycle in action by clicking on the figure in CengageNOW.

Learn more about photosynthesis in plants by clicking on the figures in CengageNOW.

9.4 (page 199) 6 Describe the flow of electrons through photosystems I and II in the noncyclic electron transport pathway and the products produced. Contrast this with cyclic electron transport. Photosystems I and II are the two types of photosynthetic units ■ involved in photosynthesis. Each photosystem includes chlorophyll

molecules and accessory pigments organized with pigment-binding proteins into antenna complexes. Only a special pair of chlorophyll a molecules in the reaction center of an antenna complex give up energized electrons to a nearby electron acceptor. P700 is in the reaction center for photosystem I; P680 is in the reaction center for photosystem II. During the noncyclic light-dependent reactions, known as noncyclic electron transport, ATP and NADPH are formed. Electrons in photosystem I are energized by the absorption of light and passed through an electron transport chain to NADP+, forming NADPH. Electrons given up by P700 in photosystem I are replaced by electrons from P680 in photosystem II. A series of redox reactions takes place as energized electrons are passed along the electron transport chain from photosystem II to photosystem I. Electrons given up by P680 in photosystem II are replaced by electrons made available by the photolysis of H2O; oxygen is released in the process. During cyclic electron transport, electrons from photosystem I are eventually returned to photosystem I. ATP is produced by chemiosmosis, but no NADPH or oxygen is generated.

9 Discuss how photorespiration reduces photosynthetic efficiency. ■

In photorespiration, C3 plants consume oxygen and generate CO2 by degrading Calvin cycle intermediates but do not produce ATP. Photorespiration is significant on bright, hot, dry days when plants close their stomata, conserving water but preventing the passage of CO2 into the leaf.

10 Compare the C4 and CAM pathways. ■

■

In the C4 pathway, the enzyme PEP carboxylase binds CO2 effectively, even when CO2 is at a low concentration. C4 reactions take place within mesophyll cells. The CO2 is fixed in oxaloacetate, which is then converted to malate. The malate moves into a bundle sheath cell, and CO2 is removed from it. The released CO2 then enters the Calvin cycle. The crassulacean acid metabolism (CAM) pathway is similar to the C4 pathway. PEP carboxylase fixes carbon at night in the mesophyll cells, and the Calvin cycle occurs during the day in the same cells. See a comparison of the C3 and C4 pathways by clicking on the figure in CengageNOW.

9.6 (page 208) 11 Contrast photoautotrophs and chemoheterotrophs with respect to their energy and carbon sources. ■ Photoautotrophs use light as an energy source and are able to incorporate atmospheric CO2 into preexisting carbon skeletons. ■ Chemoheterotrophs obtain energy by oxidizing chemicals and obtain carbon as organic molecules from other organisms.

■

Photosynthesis is the ultimate source of all chemical energy and organic molecules available to photoautotrophs, such as plants, and to virtually all other organisms as well. It also constantly replenishes the supply of oxygen in the atmosphere, vital to all aerobic organisms.

Summary Reactions for Photosynthesis The light-dependent reactions (noncyclic electron transport):

¡ Chlorophyll

6 O2 + 12 NADPH + 18 ATP The carbon fixation reactions (Calvin cycle): 12 NADPH + 18 ATP + 6 CO2 ¡ C6H12O6 + 12 NADP+ + 18 ADP + 18 Pi + 6 H2O By canceling out the common items on opposite sides of the arrows in these two coupled equations, we obtain the simplified overall equation for photosynthesis:

9.7 (page 209)

6 CO2 + H2O

12 State the importance of photosynthesis both in a plant and to other

Carbon dioxide

organisms.

Light

12 H2O + 12 NADP+ + 18 ADP + 18 Pi

Water

Light energy

¡ C6H12O6 + O2 + 6 H2O

Chlorophyll

Glucose Oxygen Water

T E S T YO U R U N D E R S TA N D I N G 1. Where is chlorophyll located in the chloroplast? (a) thylakoid membranes (b) stroma (c) matrix (d) thylakoid lumen (e) between the inner and outer membranes 2. In photolysis, some of the energy captured by chlorophyll is used to split (a) CO2 (b) ATP (c) NADPH (d) H2O (e) both b and c 3. Light is composed of particles of energy called (a) carotenoids (b) reaction centers (c) photons (d) antenna complexes (e) photosystems 4. The relative effectiveness of different wavelengths of light in photosynthesis is demonstrated by (a) an action spectrum (b) photolysis (c) carbon fixation reactions (d) photoheterotrophs (e) an absorption spectrum 5. In plants, the final electron acceptor in noncyclic electron flow is (a) NADP+ (b) CO2 (c) H2O (d) O2 (e) G3P 6. In , electrons that have been energized by light contribute their energy to add phosphate to ADP, producing ATP. (a) crassulacean acid metabolism (b) the Calvin cycle (c) photorespiration (d) C4 pathways (e) photophosphorylation 7. The mechanism by which electron transport is coupled to ATP production by means of a proton gradient is called (a) chemiosmosis (b) crassulacean acid metabolism (c) fluorescence (d) the C3 pathway (e) the C4 pathway 8. The Calvin cycle begins when CO2 reacts with (a) phosphoenolpyruvate (b) glyceraldehyde-3-phosphate (c) ribulose bisphosphate (d) oxaloacetate (e) phosphoglycerate 9. The enzyme directly responsible for almost all carbon fixation on Earth is (a) rubisco (b) PEP carboxylase (c) ATP synthase (d) phosphofructokinase (e) ligase

10. In C4 plants, C4 and C3 pathways occur at different ; whereas in CAM plants, CAM and C3 pathways . (a) times of day; locations occur at different within the leaf (b) seasons; locations (c) locations; times of day (d) locations; seasons (e) times of day; seasons 11. An organism characterized as a photoautotroph obtains energy and carbon from . from (a) light; organic molecules (b) light; CO2 (c) organic molecules; organic molecules (d) organic molecules; CO2 (e) O2; CO2 12. Label the figure. Use Figure 9-8 to check your answers.

CRITICAL THINKING

2. Only some plant cells have chloroplasts, but all actively metabolizing plant cells have mitochondria. Why? 3. Explain why the proton gradient formed during chemiosmosis represents a state of low entropy. (You may wish to refer to the discussion of entropy in Chapter 7.) 4. The electrons in glucose have relatively high free energies. How did they become so energetic? 5. SCIENCE, TECHNOLOGY, AND SOCIETY. What strategies may be employed in the future to increase world food supply? Base your answer on your knowledge of photosynthesis and related processes. 6. What would life be like for photoautotrophs if there were no chemoheterotrophs? For chemoheterotrophs if there were no photoautotrophs? 7. What might you suspect if scientists learned that a distant planet has an atmosphere that is 15% molecular oxygen? 8. EVOLUTION LINK. Propose an explanation for the fact that bacteria, chloroplasts, and mitochondria all have ATP synthase complexes.

9. ANALYZING DATA. The figure depicts the absorption spectrum of a plant pigment. What colors/wavelengths does it absorb? What is the color of this pigment?

100 Estimated absorption (%)

1. Must all autotrophs use light energy? Explain.

80 60 40 20

400

500 600 Wavelength (nm)

700

Additional questions are available in CengageNOW at www.cengage.com/ login.

Chromosomes, Mitosis, and Meiosis

Alexey Khodjakov, Wadsworth Center, Albany, NY

10

KEY CONCEPTS

Fluorescence LM of a cultured newt lung cell in mitosis (early prometaphase). The nuclear envelope has broken down, and the microtubules of the mitotic spindle (green) now interact with the chromosomes (blue).

P

re-existing cells divide to form new cells. This remarkable process enables an organism to grow, repair damaged parts, and reproduce.

10.1 In eukaryotic cells, DNA is wound around specific

Cells serve as the essential link between generations. Even the simplest

proteins to form chromatin, which in turn is folded and packaged to make individual chromosomes.

cell contains a large amount of precisely coded genetic information in

10.2 In mitosis, duplicated chromosomes separate (split

nized into informational units called genes, which control the activities

apart) and are evenly distributed into two daughter nuclei. Mitosis is an important part of the cell cycle, which consists of the successive stages through which a cell passes.

the form of deoxyribonucleic acid (DNA). An individual’s DNA is orgaof the cell and are passed on to its descendants. When a cell divides, the information contained in the DNA must be faithfully replicated and the

10.3 An internal genetic program interacts with external

copies then transmitted to each daughter cell through a precisely choreo-

signals to regulate the cell cycle.

graphed series of steps (see photograph).

10.4 Meiosis, which reduces the number of chromosome

DNA is a very long, thin molecule that could easily become tangled

sets from diploid to haploid, is necessary to maintain the normal chromosome number when two cells join during sexual reproduction. Meiosis helps to increase genetic variation among offspring.

and broken, and a eukaryotic cell’s nucleus contains a huge amount of

10.5 Meiosis and gamete production precede fertilization

structure called a chromosome, each of which contains hundreds or thou-

in the life cycles of sexually reproducing organisms.

sands of genes.

DNA. In this chapter we consider how eukaryotes accommodate the genetic material by packaging each DNA molecule with proteins to form a

We then consider mitosis, the highly regimented process that ensures a parent cell transmits one copy of every chromosome to each of its two daughter cells. In this way, the chromosome number is preserved through successive mitotic divisions. Most somatic cells (body cells) of eukaryotes divide by mitosis. Mitosis is an active area of biological research, and for good reason: errors in mitosis can result in a host of disorders and diseases

such as cancer, a disease condition in which cells divide at an uncontrolled rate and become invasive. Thus, a clearer understanding of mitosis has the potential to improve our treatment of many diseases. Finally, we discuss meiosis, a process that reduces the chromosome number by half. Sexual life cycles in eukaryotes require meiosis. Sexual reproduction involves the fusion of two sex cells, or Jan Hinsch/Photo Researchers, Inc.

gametes, to form a fertilized egg called a zygote. Meiosis makes it possible for each gamete to contain only half the number of chromosomes in the parent cell, thereby preventing the zygotes from having twice as many chromosomes as the parents.

10.1 EUKARYOTIC CHROMOSOMES ■ ■ LEARNING OBJECTIVES 1 2

Discuss the significance of chromosomes in terms of their information content. Explain how DNA is packed into chromosomes in eukaryotic cells.

The major carriers of genetic information in eukaryotes are the chromosomes, which lie within the cell nucleus. Although chromosome means “colored body,” chromosomes are virtually colorless; the term refers to their ability to be stained by certain dyes. In the 1880s, light microscopes had been improved to the point that scientists such as the German biologist Walther Fleming began to observe chromosomes during cell division. In 1903, American biologist Walter Sutton and German biologist Theodor Boveri noted independently that chromosomes were the physical carriers of genes, the genetic factors Gregor Mendel discovered in the 19th century (discussed in Chapter 11). Chromosomes are made of chromatin, a material consisting of DNA and associated proteins. When a cell is not dividing, the chromosomes are present but in an extended, partially unraveled form. Chromatin consists of long, thin threads that are somewhat aggregated, giving them a granular appearance when viewed with the electron microscope (see Fig. 4-13). During cell division, the chromatin fibers condense and the chromosomes become visible as distinct structures (FIG. 10-1).

DNA is organized into informational units called genes An organism may have thousands of genes. For example, humans have about 20,000 genes that code for proteins. As you will see in later chapters, the concept of the gene has changed considerably since the science of genetics began, but our definitions have always centered on the gene as an informational unit. By providing information needed to carry out one or more specific cell functions, a gene affects some characteristic of the organism. For example, genes govern eye color in humans, wing length in flies, and seed color in peas.

10 μm

FIGURE 10-1 Chromosomes Human chromosomes from an unidentified cell are shown in this fluorescence LM.

DNA is packaged in a highly organized way in chromosomes Prokaryotic and eukaryotic cells differ markedly in their DNA content as well as in the organization of DNA molecules. The bacterium Escherichia coli normally contains about 4 × 106 base pairs (almost 1.35 mm) of DNA in its single, circular DNA molecule. In fact, the total length of its DNA is about 1000 times as long as the length of the cell itself. Therefore, the DNA molecule is, with the help of proteins, twisted and folded compactly to fit inside the bacterial cell (see Fig. 25-2). A typical eukaryotic cell contains much more DNA than a bacterium does, and it is organized in the nucleus as multiple chromosomes; these vary widely in size and number among different species. Although a human nucleus is about the size of a large bacterial cell, it contains more than 1000 times the amount of DNA found in E. coli. The DNA content of a human sperm cell is about 3 × 109 base pairs; stretched end to end, it would measure almost 1 m long. Remarkably, this DNA fits into a nucleus with a diameter of only 10 mm. How does a eukaryotic cell pack its DNA into the chromosomes? Chromosome packaging is facilitated by certain proteins known as histones.1 Histones have a positive charge because they have a high proportion of amino acids with basic side chains (see Chapter 3). The positively charged histones associate with DNA, which has a negative charge because of its phosphate groups, to form structures called nucleosomes. The fundamental unit of each nucleosome consists of a beadlike structure with 146 base pairs of DNA wrapped around a disc-shaped core of eight histone molecules (two each of four different histone types) (FIG. 10-2).

1

A few types of eukaryotic cells lack histones. Conversely, histones occur in one group of prokaryotes, the archaea (see Chapter 25).

DNA wound around a cluster of histone molecules

Histone tails

D. E. Olins and A. L. Olins

Linker DNA

Nucleosome (10 nm diameter)

100 nm

(a) A model for the structure of a nucleosome. Each nucleosome bead contains a set of eight histone molecules, forming a protein core around which the double-stranded DNA winds. The DNA surrounding the histones consists of 146 nucleotide pairs; another segment of DNA, about 60 nucleotide pairs long, links nucleosome beads.

(b) TEM of nucleosomes from the nucleus of a chicken cell. Normally, nucleosomes are packed more closely together, but the preparation procedure has spread them apart, revealing the DNA linkers.

FIGURE 10-2 Nucleosomes

Courtesy of U. Laemmli, from Cell 12:817, 1988. Copyright by Cell Press

Although the nucleosome was originally defined as a bead plus a DNA segment that links it to an adjacent bead, today the term more commonly refers only to the bead itself (that is, the eight histones and the DNA wrapped around them). Nucleosomes function like tiny spools, preventing DNA from becoming tangled. You can see the importance of this role in FIGURE 10-3, which illustrates the enormous length of DNA that unravels from a mouse chromosome after researchers have removed the histones. The role of histones is more than simply structural because their arrangement also affects the activity of the DNA with which they are associated. Histones are increasingly viewed as an important part of the regulation of gene expression—that is, whether genes are turned off or on. We discuss gene regulation by histones in Chapter 14. The wrapping of DNA into nucleosomes represents the first level of chromosome structure. FIGURE 10-4 shows the higher-order structures of chromatin leading to the formation of a condensed chromosome. The nucleosomes themselves are 10 nm in diameter. The packed nucleosome state occurs when a fifth type of histone, known as histone H1, associates with the linker DNA, packing adjacent nucleosomes together to form a compacted 30 nm chromatin fiber. In extended chromatin, these fibers form large, coiled loops held together by scaffolding proteins, nonhistone proteins that help maintain chromosome structure. The loops then interact to form the condensed chromatin found in a chromosome. Cell biologists have identified a group of proteins, collectively called condensin, required for chromosome compaction. Condensin binds to DNA and wraps it into coiled loops that are compacted into a mitotic or meiotic chromosome.

DNA loops

Scaffolding proteins

2 μm

FIGURE 10-3 TEM of a mouse chromosome depleted of histones Notice how densely packed the DNA strands are, even though they have been released from the histone proteins that organize them into tightly coiled structures.

Chromosome number and informational content differ among species Every individual of a given species has a characteristic number of chromosomes in the nuclei of its somatic (body) cells. However, it is not the number of chromosomes that makes each species unique but

KEY POINT

When a cell prepares to divide, its chromosomes become thicker and shorter as their long chromatin fibers are compacted. 1400 nm

K. G. Murti/Visuals Unlimited. Inc.

700 nm

300 nm fiber (looped domains)

30 nm chromatin fiber

Condensed chromosome Condensed chromatin

DNA wound around a cluster of histone molecules

Scaffolding protein Extended chromatin

Packed nucleosomes

Histone 10 nm Nucleosomes

2 nm DNA double helix

FIGURE 10-4 Animated Organization of a eukaryotic chromosome This diagram shows how DNA is packaged into highly condensed metaphase chromosomes. First, DNA is wrapped around histone proteins to form nucleosomes. Then, the nucleosomes are compacted into chroma-

the information the genes specify. Most human somatic cells have exactly 46 chromosomes, but humans are not humans merely because we have 46 chromosomes. Some other species—the olive tree, for example—also have 46. Some humans have an abnormal chromosome composition with more or fewer than 46 (see Fig. 16-4). Other species have different chromosome numbers. A certain species of roundworm has only 2 chromosomes in each cell, whereas some crabs have as many as 200, and some ferns have more than 1000. Most animal and plant species have between 8 and 50 chromosomes per somatic cell. Numbers above and below

tin fibers, which are coiled into looped domains. The looped domains are compacted, ultimately forming chromosomes.

these are uncommon. The number of chromosomes a species has does not indicate the species’ complexity or its status within a particular domain or kingdom.

Review ■

■

What are the informational units on chromosomes called? Of what do these informational units consist? How is the large discrepancy between DNA length and nucleus size addressed in eukaryotic cells?

10.2 THE CELL CYCLE AND MITOSIS

KEY POINT

PH INTER ASE

■ ■ LEARNING OBJECTIVES 3 4 5

Identify the stages in the eukaryotic cell cycle and describe their principal events. Describe the structure of a duplicated chromosome, including the sister chromatids, centromeres, and kinetochores. Explain the significance of mitosis and describe the process.

When cells reach a certain size, they usually either stop growing or divide. Not all cells divide; some, such as skeletal muscle and red blood cells, do not normally divide once they are mature. Other cells undergo a sequence of activities required for growth and cell division. The stages through which a cell passes from one cell division to the next are collectively referred to as the cell cycle. Timing of the cell cycle varies widely, but in actively growing plant and animal cells, it is about 8 to 20 hours. The cell cycle consists of two main phases, interphase and M phase, both of which can be distinguished under a light microscope (FIG. 10-5).

Chromosomes duplicate during interphase Most of a cell’s life is spent in interphase, the time when no cell division is occurring. A cell is metabolically active during interphase, synthesizing needed materials (proteins, lipids, and other biologically important molecules) and growing. Here is the sequence of interphase and M phase in the eukaryotic cell cycle: G1 phase

S phase Interphase

G2 phase

The cell cycle is the successive series of events in the life of a cell.

G1 (First gap phase)

S (Synthesis phase)

G2 (Second gap phase)

M PHASE (Mitosis and cytokinesis)

FIGURE 10-5 Animated The eukaryotic cell cycle The cell cycle includes interphase (G1, S, and G2) and M phase (mitosis and cytokinesis). Proportionate amounts of time spent at each stage vary among species, cell types, and growth conditions. If the cell cycle were a period of 12 hours, G1 would be about 5 hours, S would be 4.5 hours, G2 would be 2 hours, and M phase would be 30 minutes.

mitosis and cytokinesis M phase

The time between the end of mitosis and the beginning of the S phase is termed the G1 phase (G stands for gap, an interval during which no DNA synthesis occurs). Growth and normal metabolism take place during the G1 phase, which is typically the longest phase. Cells that are not dividing usually become arrested in this part of the cell cycle and are said to be in a state called G0. Toward the end of G1, the enzymes required for DNA synthesis become more active. Synthesis of these enzymes, along with proteins needed to initiate cell division (discussed later in the chapter), enable the cell to enter the S phase. During the synthesis phase, or S phase, DNA replicates and histone proteins are synthesized so that the cell can make duplicate copies of its chromosomes. How did researchers identify the S phase of the cell cycle? In the early 1950s, researchers demonstrated that cells preparing to divide duplicate their chromosomes at a relatively restricted time interval during interphase and not during early mitosis, as previously hypothesized. These investigators used isotopes, such as 3H, to synthesize radioactive thymidine, a nucleotide that is incorporated specifically into DNA as it is synthesized. After radioactive thymidine was supplied for a

brief period (such as 30 minutes) to actively growing cells, autoradiography (see Fig. 2-3) on exposed film showed that a fraction of the cells had silver grains over their chromosomes. The nuclei of these cells were radioactive because during the experiment DNA had replicated. DNA replication was not occurring in the cells that did not have radioactively labeled chromosomes. Researchers therefore inferred that the proportion of labeled cells out of the total number of cells provides a rough estimate of the length of the S phase relative to the rest of the cell cycle. After it completes the S phase, the cell enters a second gap phase, the G2 phase. At this time, increased protein synthesis occurs, as the final steps in the cell’s preparation for division take place. For many cells, the G2 phase is short relative to the G1 and S phases. M phase involves two main processes, mitosis and cytokinesis. Mitosis, the nuclear division that produces two nuclei containing chromosomes identical to the parental nucleus, begins at the end of the G2 phase. Cytokinesis, which generally begins before mitosis is complete, is the division of the cell cytoplasm to form two cells. Mitosis is a continuous process, but for descriptive purposes, it is divided into five stages: prophase ¡ prometaphase ¡ metaphase ¡ anaphase ¡ telophase

PROPHASE

PROMETAPHASE

Phototake; Prometaphase: Ed Reschke/Peter Arnold, Inc.

INTERPHASE

Chromatin

Nucleolus Nucleus Fragments of nuclear envelope

Sister chromatids of duplicated chromosome

Kinetochore Spindle microtubule

Nuclear envelope Centrioles

Developing mitotic spindle

Plasma membrane

(a) Cell carries out normal life activities. Chromosomes become duplicated.

(b) Long fibers of chromatin condense as compact mitotic chromosomes, each consisting of two chromatids attached at their centromeres. Cytoskeleton is disassembled, and mitotic spindle forms between centrioles, which have moved to poles of cell. Nuclear envelope begins to disassemble.

(c) Spindle microtubules attach to kinetochores of chromosomes. Chromosomes begin to move toward cell’s midplane.

FIGURE 10-6 Animated Interphase and the stages of mitosis The LMs show plant cells, which lack centrioles. The drawings depict generalized animal cells with a diploid chromosome number of four; the sizes of the nuclei and chromosomes are exaggerated to show the structures more clearly.

Study FIGURE 10-6 while you read the following descriptions of these stages as they occur in a typical plant or animal cell.

During prophase, duplicated chromosomes become visible with the microscope The first stage of mitosis, prophase, begins with chromosome compaction, when the long chromatin fibers that make up the chromosomes begin a coiling process that makes them shorter and thicker. The compacted chromatin can then be distributed to the daughter cells with less likelihood of tangling. When stained with certain dyes and viewed through the light microscope, chromosomes become visible as darkly stained bodies as prophase progresses. It is now apparent that each chromosome was duplicated during the preceding S phase and consists of a pair of sister chromatids, which contain identical, double-stranded DNA sequences. Each chromatid includes a constricted region called the centromere. Sister chromatids are tightly associated in the vicinity of their centromeres (FIG. 10-7). The chemical basis for

this close association at the centromeres consists of precise DNA sequences that are tightly bound to specific proteins. For example, sister chromatids are physically linked by a ringshaped protein complex called cohesin. Cohesins extend along the length of the sister chromatid arms and are particularly concentrated at the centromere (FIG. 10-8). These cohesins, which hold the replicated chromosomes together from their synthesis in S phase onward, help to ensure accurate chromosome segregation during mitosis. Attached to each centromere is a kinetochore, a multiprotein complex to which microtubules can bind. These microtubules function in chromosome distribution during mitosis, in which one copy of each chromosome is delivered to each daughter cell. A dividing cell can be described as a globe, with an equator that determines the midplane (equatorial plane) and two opposite poles. This terminology is used for all cells regardless of their actual shape. Microtubules radiate from each pole, and some of these protein fibers elongate toward the chromosomes, forming the mitotic spindle, a structure that separates the duplicated chromosomes

ANAPHASE

TELOPHASE

25 μm

METAPHASE

Spindle Cleavage furrow

Centriole pair at spindle pole Cell’s midplane (metaphase plate)

(d) Chromosomes line up along cell’s midplane. Spindle microtubules attach each chromosome to both poles.

Daughter chromosomes

(e) Sister chromatids separate at their centromeres. One group of chromosomes moves toward each pole of cell. Spindle poles move farther apart.

Re-forming nuclear envelope

(f) Chromosomes are grouped at poles. Chromosomes decondense, and nuclear envelopes begin to form. Cytokinesis produces two daughter cells.

▲

▲ FIGURE 10-6 Continued

FIGURE 10-7 Sister chromatids and centromeres

Centromere region Microtubules

Kinetochore

E. J. DuPraw

Sister chromatids

1.0 μm

The sister chromatids, each consisting of tightly coiled chromatin fibers, are tightly associated at their centromere regions, indicated by the brackets. Associated with each centromere is a kinetochore, which serves as a microtubule attachment site. Kinetochores and microtubules are not visible in this TEM of a metaphase chromosome.

during anaphase (FIG. 10-9). The minus ends of these microtubules are at the poles, and the plus ends extend to the cell’s midplane. It might be helpful to review Figure 4-24, which shows the organization of microtubules as linear polymers of the protein tubulin. The organization and function of the spindle require the presence of motor proteins and a variety of signaling molecules. Animal cells differ from plant cells in the details of mitotic spindle formation. In both types of dividing cells, each pole contains a region, the microtubule-organizing center, from which extend the microtubules that form the mitotic spindle. The electron microscope shows that microtubule-organizing centers in certain plant cells consist of fibrils with little or no discernible structure. In contrast, animal cells have a pair of centrioles in the middle of each microtubule-organizing center (see Fig. 4-26). The centrioles are surrounded by fibrils that make up the pericentriolar

KEY POINT

When chromosomes duplicate, sister chromatids are initially linked to one another by protein complexes called cohesins. Cohesin linkages are particularly concentrated in the vicinity of the centromere. Sister chromatids of duplicated chromosome Kinetochore

Prophase

1

Cohesin complexes

As mitosis progresses, cohesins dissociate from the duplicated chromosome arms.

Cohesin complexes in centromere region

Spindle microtubule Metaphase

2

Cohesins then dissociate at the centromere, allowing the daughter chromosomes to separate during anaphase.

Daughter chromosomes

Anaphase

FIGURE 10-8 Cohesins

material. The spindle microtubules terminate in the pericentriolar material, but they do not actually touch the centrioles. Although cell biologists once thought spindle formation in animal cells required centrioles, their involvement is probably coincidental. Current evidence suggests that centrioles organize the pericentriolar material and ensure its duplication when the centrioles duplicate.

Each of the two centrioles is duplicated during the S phase of interphase, yielding two centriole pairs. Late in prophase, microtubules radiate from the pericentriolar material surrounding the centrioles; these clusters of microtubules are called asters. The two asters migrate to opposite sides of the nucleus, establishing the two poles of the mitotic spindle.

Prometaphase begins when the nuclear envelope breaks down During prometaphase the nuclear envelope fragments so that the spindle microtubules come into contact with the chromosomes; units of the disassembled nuclear envelope are sequestered in vesicles to be used later, to assemble nuclear envelopes for the daughter cells. The nucleolus shrinks and usually disappears, and the mitotic spindle is completely assembled. At the start of prometaphase, the duplicated chromosomes are scattered throughout the nuclear region (see the chapter introduction photograph, which shows early prometaphase). The spindle microtubules grow and shrink as they move toward the center of the cell in a “search and capture” process. Their random, dynamic movements give the appearance that they are “searching” for the chromosomes. If a microtubule comes near a centromere, one of the kinetochores of a duplicated chromosome “captures” it. As the now-tethered chromosome continues moving toward the cell’s midplane, the unattached kinetochore of its sister chromatid becomes connected to a spindle microtubule from the cell’s other pole. During the chromosomes’ movements toward the cell’s midplane, long microtubules are shortened by the removal of tubulin subunits, and the short microtubules are lengthened by the addition of tubulin subunits. Evidence indicates that shortening and lengthening occurs at the kinetochore end (the plus end) of the microtubule, not at the spindle pole end (the minus end). This shortening or lengthening occurs while the spindle microtubule remains firmly tethered to the kinetochore. Motor proteins located at the kinetochores may be involved in this tethering of the spindle microtubule. These motor proteins may work in a similar fashion to the kinesin motor shown in Figure 4-25. To summarize the events of prometaphase, sister chromatids of each duplicated chromosome become attached at their kinetochores to spindle microtubules extending from opposite poles of the cell, and the chromosomes begin to move toward the cell’s midplane. As the cell transitions from prometaphase to metaphase, cohesins dissociate from the sister chromatid arms, freeing them from one another, although some cohesins remain in the vicinity of the centromere.

Duplicated chromosomes line up on the midplane during metaphase During metaphase, all the cell’s chromosomes align at the cell’s midplane, or metaphase plate. As already mentioned, one of the two sister chromatids of each chromosome is attached by its kinetochore to microtubules from one pole, and its sister chromatid is attached by its kinetochore to microtubules from the opposite pole. The mitotic spindle has three types of microtubules: polar microtubules, kinetochore microtubules, and astral microtubules

Metaphase plate (cell’s midplane) Kinetochore microtubule (chromosome spindle fiber)

Centrioles Astral microtubules Pericentriolar material

Sister chromatids

(a) One end of each microtubule of this animal cell is associated with one of the poles. Astral microtubules (green) radiate in all directions, forming the aster. Kinetochore microtubules (red ) connect the kinetochores to the poles, and polar (nonkinetochore) microtubules (blue) overlap at the midplane.

CNRI/Phototake, NYC

Polar (nonkinetochore) microtubule

10 μm

(b) This fluorescence LM of an animal cell at metaphase shows a well-defined spindle and asters (chromosomes, orange; microtubules, green).

FIGURE 10-9 The mitotic spindle

(see Fig. 10-9). Polar microtubules, also known as nonkinetochore microtubules, extend from each pole to the equatorial region, where they overlap and interact with nonkinetochore microtubules from the opposite pole. Kinetochore microtubules extend from each pole and attach to chromosomes at their kinetochores. Astral microtubules are the short microtubules that form asters at each pole. Each chromatid is completely condensed and appears distinct during metaphase. Because individual chromosomes are more obvious at metaphase than at any other time, the karyotype, or chromosome composition, is usually checked at this stage for chromosome abnormalities (see Chapter 16). As the mitotic cell transitions from metaphase to anaphase, the remaining cohesin proteins joining the sister chromatids at their centromere dissociate.

During anaphase, chromosomes move toward the poles Anaphase begins as the sister chromatids separate. Once the chromatids are no longer attached to their duplicates, each chromatid is called a chromosome. The now-separate chromosomes move to opposite poles, using the spindle microtubules as tracks. The kinetochores, still attached to kinetochore microtubules, lead the way, with the chromosome arms trailing behind. Anaphase ends when all the chromosomes have reached the poles. Cell biologists are making significant progress in understanding the overall mechanism of chromosome movement in anaphase. Chromosome movements are studied in several ways. The

number of microtubules at a particular stage or after certain treatments is determined by carefully analyzing electron micrographs. Researchers also physically perturb living cells that are dividing, using microlaser beams or mechanical devices known as micromanipulators. A skilled researcher can move chromosomes, break their connections to microtubules, and even remove them from the cell entirely. Microtubules lack elastic or contractile properties. Then how do the chromosomes move apart? Are they pushed or pulled, or do other forces operate? Microtubules are dynamic structures, with tubulin subunits constantly being removed from their ends and others being added. Evidence indicates that during anaphase, kinetochore microtubules shorten, or depolymerize, at their plus ends, that is, closest toward the midplane of the cell (FIG. 10-10). This shortening mechanism pulls the chromosomes toward the poles. A second mechanism also plays a role in chromosome separation. During anaphase the spindle as a whole elongates, at least partly because polar microtubules originating at opposite poles are associated with motors that let them slide past one another at the midplane. The sliding decreases the degree of overlap, thereby “pushing” the poles apart. This mechanism indirectly causes the chromosomes to move apart because they are attached to the poles by kinetochore microtubules.

During telophase, two separate nuclei form During the final stage of mitosis, telophase, chromosomes arrive at the poles and there is a return to interphase-like conditions. The

KEY EXPERIMENT QUESTION: Do spindle microtubules move chromosomes by a shortening (that is, depolymerization) of microtubules at the spindle poles or at the kinetochore ends?

HYPOTHESIS: Spindle microtubules move chromosomes toward the spindle poles by a mechanism in which the spindle microtubules are shortened at their kinetochore ends. EXPERIMENT: Microtubules in pig kidney cells in early anaphase were labeled with a fluorescent dye that specifically attaches to microtubules.

The researchers marked the microtubules by using a laser microbeam to bleach the dye while keeping the microtubules intact.

Cytokinesis forms two separate daughter cells Cytokinesis, the division of the cytoplasm to yield two daughter cells, is the last step in M phase and usually overlaps mitosis, generally beginning during telophase. Cytokinesis of an animal or fungal cell (yeast, for example) begins as an actomyosin contractile ring is assembled and attached to the plasma membrane. The contractile ring encircles the cell in the equatorial region, at right angles to the spindle (FIG. 10-11a). The contractile ring consists of an association between actin and myosin filaments; it is thought that the motor activity of myosin moves actin filaments to cause the constriction, similar to the way actin and myosin cause muscle contraction (see Fig. 40-10). The ring contracts, producing a cleavage furrow that gradually deepens and eventually separates the cytoplasm into two daughter cells, each with a complete nucleus. The contractile ring then disassembles. In plant cells, cytokinesis occurs by forming a cell plate (FIG. 10-11b), a partition constructed in the equatorial region of the spindle that grows laterally toward the cell wall. The cell plate forms as a line of vesicles originating in the Golgi complex. The vesicles contain materials to construct both a primary cell wall for each daughter cell and a middle lamella that cements the primary cell walls together. The vesicle membranes fuse to become the plasma membrane of each daughter cell. Multinucleated cells form if mitosis is not followed by cytokinesis; this is a normal condition for certain cell types. For example, the body of plasmodial slime molds consists of a multinucleate mass of cytoplasm (see Fig. 26-19a).

Mitosis produces two cells genetically identical to the parent cell RESULTS AND CONCLUSION: The chromosomes moved toward the bleached areas of the spindle microtubules, indicating a shortening of the microtubules on the kinetochore side. The microtubules on the polar ends did not shorten.

Chromosomes moved poleward because they remained anchored to the kinetochore microtubules as tubulin subunits were removed at the kinetochore ends of the microtubules.

FIGURE 10-10 Animated Using laser photobleaching to determine how chromosomes are transported toward the spindle poles during anaphase

chromosomes decondense by partially uncoiling. A new nuclear envelope forms around each set of chromosomes, made at least in part from small vesicles and other components derived from the old nuclear envelope. The spindle microtubules disappear, and the nucleoli reorganize.

The remarkable regularity of the process of cell division ensures that each daughter nucleus receives exactly the same number and kinds of chromosomes that the parent cell had. Thus, with a few exceptions, every cell of a multicellular organism has the same genetic makeup. If a cell receives more or fewer than the characteristic number of chromosomes through some malfunction of the cell division process, the resulting cell may show marked abnormalities and often cannot survive. Mitosis provides for the orderly distribution of chromosomes (and of centrioles, if present), but what about the various cytoplasmic organelles? For example, all eukaryotic cells, including plant cells, require mitochondria. Likewise, photosynthetic plant cells cannot carry out photosynthesis without chloroplasts. These organelles contain their own DNA and appear to form by the division of previously existing mitochondria or plastids or their precursors. This nonmitotic division process is similar to prokaryotic cell division (discussed in the next section) and generally occurs during interphase. Because many copies of each organelle are present in each cell, organelles are apportioned with the cytoplasm that each daughter cell receives during cytokinesis.

Lacking nuclei, prokaryotes divide by binary fission Bacteria and archaea contain much less DNA than do most eukaryotic cells, but precise distribution of the genetic material into two

daughter cells is still a formidable process. Prokaryotic DNA usually consists of a single, circular chromosome that is packaged with associated proteins. Although the distribution of genetic material in dividing prokaryotic cells is a simpler process than mitosis, it nevertheless is very precise, to ensure that the daughter cells are genetically identical to the parent cell. Prokaryotes reproduce asexually, generally by binary fission, a process in which one cell divides into two daughter cells (FIG. 10-12). The circular DNA molecule replicates, resulting in two identical chromosomes. DNA replication begins at a single site on the bacterial chromosome, termed the origin of replication. DNA synthesis proceeds from that point in both directions until they eventually meet (see Fig. 12-16a). Following replication, the daughter chromosomes separate and move to opposite ends of the elongating cell. Cytokinesis between the daughter chromosomes is controlled by the Z ring, a

protein scaffold that holds about 10 different proteins around the cell’s midsection. There the plasma membrane grows inward between the two DNA copies, dividing the cell’s cytoplasm in half, and a new transverse cell wall is synthesized between the two cells. (Bacterial reproduction is described further in Chapter 25.)

Review ■

■

What are the stages of the cell cycle? During which stage does DNA replicate? What are the stages of mitosis and what happens in each stage?

10.3 REGULATION OF THE CELL CYCLE ■ ■ LEARNING OBJECTIVE 6 Explain some ways in which the cell cycle is controlled.

When conditions are optimal, some prokaryotic cells can divide every 20 minutes. The generation times of eukaryotic cells are generally much longer, although the frequency of cell division varies widely among different species and among different tissues of the same species. Some skeletal muscle cells usually stop dividing after the first few months of life, whereas blood-forming cells, digestive tract cells, and skin cells divide frequently Actomyosin throughout the life of the organism. Under optimal concontractile ring ditions of nutrition, temperature, and pH, the length of the eukaryotic cell cycle is constant for any given cell type. Under less favorable conditions, however, the generation time may be longer. Certain regulatory molecules that control the cell cycle are common to all eukaryotes. Genetically programmed in the cell’s

10 μm

(a) TEM of the equatorial region of a cultured animal cell undergoing cytokinesis. Note the cleavage furrow. Dividing fungal cells also have a contractile ring that causes cytokinesis. Vesicles gather on cell's midplane

Plasma membrane

Cell wall

Small vesicles fuse, forming larger vesicles

Eventually one large vesicle exists

Cell plate forming

(b) Cell plate formation during cytokinesis in a plant cell. The TEM shows a maple leaf cell (Acer saccharinum ) undergoing cytokinesis.

FIGURE 10-11 Animated Cytokinesis in animal and plant cells The nuclei in both TEMs are in telophase. The drawings show 3-D relationships.

Nucleus

Cell plate forming

New cell walls (from vesicle contents)

New plasma membranes (from vesicle membranes)

E. H. Newcomb and B. A. Palevitz/Biological Photo Service

T. E. Schroeder/Biological Photo Service

Cleavage furrow

5 μm

The resulting daughter cells might have too few or too many chromosomes. An abnormal number of chromosomes is associated with Down syndrome as well as Bacterial cell many cancers. 1 DNA replication begins Plasma FIGURE 10-14 shows some of the membrane at single site on bacterial Bacterial key molecules involved in regulating chromosome. Cell wall chromosome the cell cycle. Among them are protein Origin of kinases, enzymes that activate or inacreplication tivate other proteins by phosphorylating (adding phosphate groups to) them. The protein kinases involved in controlling Unreplicated the cell cycle are cyclin-dependent kiregion of 2 Replication continues, as nases (Cdks). The activity of various chromosome replication enzymes work in Two copies Cdks increases and then decreases as the both directions from site of bacterial cell moves through the cell cycle. Cdks DNA where replication began. are active only when they bind tightly to regulatory proteins called cyclins. The cyclins are so named because their levels Z ring fluctuate predictably during the cell cycle (that is, they “cycle,” or are alternately 3 Replication is completed, synthesized and degraded as part of the and daughter chromosomes separate. Z ring forms at cell’s cell cycle). midsection. Cell begins to Three scientists who began their divide, as plasma membrane research in the 1970s and 1980s on the grows inward. A new cell wall roles of protein kinases and cyclins in New cell forms. walls the cell cycle (Leland Hartwell from the United States, and Paul Nurse and Tim 4 Binary fission is complete. Hunt from Great Britain) were awarded Two identical daughter cells result. the Nobel Prize in Physiology or Medicine in 2001. Their discoveries were cited as important not only in working out the Two identical bacterial cells details of the fundamental cell process of mitosis but also in understanding why cancer cells divide when they should not. FIGURE 10-12 Binary fission For example, cyclin levels are often higher The bacterial chromosome is much longer than depicted here and is tethered to the plasma membrane at one spot (not shown). than normal in human cancer cells. When a specific Cdk associates with a specific cyclin, it forms a cyclin–Cdk nucleus, these regulatory molecules are components of the cell complex. Cyclin–Cdk complexes phosphorylate enzymes and cycle control system found in organisms as diverse as yeast, clams, other proteins. Some of these proteins become activated when they frogs, humans, and plants. Regulatory molecules trigger a specific are phosphorylated, and others become inactivated. For example, sequence of events during the cell cycle. phosphorylation of the protein p27, known to be a major inhibitor Because the cell cycle consists of hundreds of sequential of cell division, is thought to initiate degradation of the protein. As events that proceed in an orderly manner, a failure to carefully various enzymes are activated or inactivated by phosphorylation, control these events can have disastrous consequences. Control the activities of the cell change. Thus, a decrease in a cell’s level of mechanisms in the genetic program, called cell-cycle checkp27 causes a nondividing cell to resume division. points, temporarily block key events from being initiated during Eukaryotic cells form four major cyclin–Cdk complexes: G1Cdk, G1/S-Cdk, S-Cdk, and M-Cdk. Each cyclin–Cdk complex the cell cycle. Cell-cycle checkpoints ensure that all the events of a phosphorylates a different group of proteins. G1-Cdk prepares the particular stage have been completed before the next stage begins cell to pass from the G1 phase to the S phase, and then G1/S-Cdk (FIG. 10-13). The checkpoints are inactivated after they have done their job so the cell cycle can proceed. commits the cell to undergo DNA replication. S-Cdk initiates DNA Genes that encode molecules involved in checkpoints are critireplication. M-Cdk promotes the events of mitosis, including chrocally important to the cell cycle. If a checkpoint gene is defective, it mosome condensation, nuclear envelope breakdown, and mitotic can result in cancer or other serious diseases. Consider what might spindle formation. happen if the metaphase-anaphase checkpoint molecules were M-Cdk also activates another enzyme complex, the anaphasenonfunctional. In this case, anaphase might be initiated too early, promoting complex (APC), toward the end of metaphase. APC before all chromosomes were properly attached to spindle fibers. initiates anaphase by allowing degradation of the cohesins and

KEY POINT

Binary fission is a precisely orchestrated sequence of events that ensures each bacterial daughter cell has identical genetic material.

KEY POINT

1

When a cell has not completed the steps leading to a cell-cycle checkpoint, the checkpoint is active and halts the cell cycle. When the necessary steps are completed, the checkpoint is inactivated, and the cell cycle proceeds.

G1-S checkpoint: The first key checkpoint ensures that the cell has the necessary growth factors, nutrients, and enzymes to synthesize DNA. Without the proper signals that the cell is ready to proceed, the checkpoint will not let DNA synthesis begin.

1

G1 (First gap phase)

3

Metaphase-anaphase checkpoint: Sometimes called the spindle checkpoint, this checkpoint occurs at the end of metaphase and prevents anaphase from occurring until all kinetochores are properly attached to spindle fibers along the cell’s midplane.

3

S (Synthesis phase)

G2 (Second gap phase) 2

2

G2-M checkpoint: This cell-cycle checkpoint ensures that DNA replication is finished before the cell begins mitosis. If a cell has damaged or unreplicated DNA, the checkpoint will not let the cell undergo mitosis.

FIGURE 10-13 Key checkpoints in the cell cycle The cell cycle consists of hundreds of sequential events. The red bars show three important checkpoints that determine that previous steps are completed so that the next steps may proceed. Each checkpoint is inactivated after it has performed its function, allowing the cell cycle to continue.

other proteins that hold the sister chromatids together during metaphase. As a result, the sister chromatids separate as two daughter chromosomes. At this point, cyclin is degraded to negligible levels and M-Cdk activity drops, allowing the mitotic spindle to disassemble and the cell to exit mitosis. Certain drugs can stop the cell cycle at a specific checkpoint. Some of these prevent DNA synthesis, whereas others inhibit the synthesis of proteins that control the cycle or inhibit the synthesis of structural proteins that contribute to the mitotic spindle. Because one of the distinguishing features of most cancer cells is their high rate of cell division relative to that of most normal somatic cells, cancer cells are greatly affected by these drugs. Many side effects of certain anticancer drugs (such as nausea and hair loss) are due to the drugs’ effects on normal cells that divide rapidly in the digestive system and hair follicles. In plant cells, certain hormones stimulate mitosis. These include the cytokinins, a group of plant hormones that promote mitosis both in normal growth and in wound healing (see Chapter 38). Similarly, animal hormones, such as certain steroids, stimulate growth and mitosis (see Chapter 49). Protein growth factors, which are active at extremely low concentrations, stimulate mitosis in some animal cells. Of the approximately 50 protein growth factors known, some act only on

specific types of cells, whereas others act on a broad range of cell types. For example, the effects of the growth factor erythropoietin are limited to cells that will develop into red blood cells, but epidermal growth factor stimulates many cell types to divide. Many types of cancer cells divide in the absence of growth factors.

Review ■ ■

What are cell-cycle checkpoints? What are two molecular controls that trigger the onset of different stages of the cell cycle?

10.4 SEXUAL REPRODUCTION AND MEIOSIS ■ ■ LEARNING OBJECTIVES 7 Differentiate between asexual and sexual reproduction. 8 Distinguish between haploid and diploid cells, and define homologous chromosomes.

9 Explain the significance of meiosis and describe the process. 10 Contrast mitosis and meiosis, emphasizing the different outcomes.

KEY POINT

Cyclin-dependent kinases (Cdks) control the phosphorylation of other proteins, thereby regulating the transitions between phases of the cell cycle. 1

Cyclin is synthesized and accumulates.

Cdk G1 5

S

Cdk is not degraded but is recycled and reused. M

G2

Cyclin 4

An activated enzyme complex recognizes a specific amino acid Degraded sequence in cyclin and targets it for cyclin destruction. When cyclin is degraded, M-Cdk activity is terminated, and the cells formed by mitosis enter G1.

M-Cdk (triggers M phase) 3

Cdk

2

Cdk associates with cyclin, forming M-Cdk, a cyclin–Cdk complex.

M-Cdk phosphorylates proteins, activating those that facilitate mitosis and inactivating those that inhibit mitosis.

FIGURE 10-14 Molecular control of the cell cycle This diagram is a simplified view of the control system that triggers the cell to move from G2 to M phase.

Although the details of the reproductive process vary greatly among different kinds of eukaryotes, biologists distinguish two basic types of reproduction: asexual and sexual. In asexual reproduction a single parent splits, buds, or fragments to produce two or more individuals. In most kinds of eukaryotic asexual reproduction, all the cells are the result of mitotic divisions, so their genes and inherited traits are like those of the parent. Such a group of genetically identical organisms is called a clone. In asexual reproduction, organisms that are well adapted to their environment produce new generations of similarly adapted organisms. Asexual reproduction occurs rapidly and efficiently, partly because the organism does not need to expend time and energy finding a mate. In contrast, sexual reproduction involves the union of two sex cells, or gametes, to form a single cell called a zygote. Usually two different parents contribute the gametes, but in some cases a single parent furnishes both gametes. In the case of animals and plants, the egg and sperm cells are the gametes, and the fertilized egg is the zygote. Sexual reproduction results in genetic variation among the offspring. (How this genetic variation arises is discussed later in this chapter and in Chapter 11.) Because the offspring produced by sexual reproduction are not genetically identical to their parents or to each other, some offspring may be able to survive environmental

changes better than either parent does. However, one disadvantage of sexual reproduction is that some offspring with a different combination of traits may be less likely to survive than their parents. There is a potential problem in eukaryotic sexual reproduction: if each gamete had the same number of chromosomes as the parent cell that produced it, then the zygote would have twice as many chromosomes. This doubling would occur generation after generation. How do organisms avoid producing zygotes with everincreasing chromosome numbers? To answer this question, we need more information about the types of chromosomes found in cells. Each chromosome in a somatic cell of a plant or animal normally has a partner chromosome. The two partners, known as homologous chromosomes, are similar in size, shape, and the position of their centromeres. Furthermore, special chromosome-staining procedures produce a characteristic pattern of bands evident in the members of each chromosome pair. In most species, chromosomes vary enough in their structure that cell biologists can distinguish the different chromosomes and match up the homologous pairs. The 46 chromosomes in human cells constitute 23 homologous pairs. The most important feature of homologous chromosomes is that they carry information about the same genetic traits, although this information is not necessarily identical. For example, each member of a homologous pair may carry a gene that specifies he-

moglobin structure. But one member may have the information for the normal hemoglobin b chain (see Fig. 3-22a), whereas the other may specify the abnormal form of hemoglobin associated with sickle cell anemia (see Chapter 16). Homologous chromosomes can therefore be contrasted with the two members of a pair of sister chromatids, which are precisely identical to each other. A set of chromosomes has one of each kind of chromosome; in other words, it contains one member of each homologous pair. If a cell or nucleus contains two sets of chromosomes, it is said to have a diploid chromosome number. If it has only a single set of chromosomes, it has the haploid number. In humans, the diploid chromosome number is 46 and the haploid number is 23. When a sperm and egg fuse at fertilization, each gamete is haploid, contributing one set of chromosomes; the diploid number is thereby restored in the fertilized egg (zygote). When the zygote divides by mitosis to form the first two cells of the embryo, each daughter cell receives the diploid number of chromosomes, and subsequent mitotic divisions repeat this. Thus, somatic cells are diploid. An individual whose cells have three or more sets of chromosomes is polyploid. Polyploidy is relatively rare among animals but common among plants (see Chapter 20). In fact, polyploidy has been an important mechanism of plant evolution. As many as 80% of all flowering plants are polyploid. Polyploid plants are often larger and hardier than diploid members of the same group. Many commercially important plants, such as wheat and cotton, are polyploid. The chromosome number found in the gametes of a particular species is represented as n, and the zygotic chromosome number is represented as 2n. If the organism is not polyploid, the haploid chromosome number is equal to n, and the diploid number is equal to 2n; thus, in humans, n = 23 and 2n = 46. For simplicity, in the rest of this chapter, the organisms used as examples are not polyploid. We use diploid and 2n interchangeably, and haploid and n interchangeably, although the terms are not strictly synonymous.

Interphase

One 2n cell with four unduplicated chromosomes (2 homologous pairs) Replication of DNA

Four chromosomes, each with two sister chromatids

Homologous pair of duplicated chromosomes

Meiosis I: Homologous chromosomes pair and then separate

Sister chromatids

Meiosis II: Sister chromatids separate

Meiosis produces haploid cells with unique gene combinations A cell division that reduces chromosome number is called meiosis. The term meiosis means “to make smaller,” and the chromosome number is reduced by one-half. In meiosis a diploid cell undergoes two cell divisions, potentially yielding four haploid cells. It is important to note that haploid cells do not contain just any combination of chromosomes, but one member of each homologous pair. The events of meiosis are similar to the events of mitosis, with four important differences:

This figure begins with a diploid cell with four unduplicated chromosomes. The chromosomes derived from one parent are shown in blue, and those from the other parent are red. Homologous pairs are similar in size and shape.

1. Meiosis involves two successive nuclear and cytoplasmic divisions, producing up to four cells. 2. Despite two successive nuclear divisions, the DNA and other chromosome components duplicate only once—during the interphase preceding the first meiotic division. 3. Each of the four cells produced by meiosis contains the haploid chromosome number, that is, only one chromosome set containing only one representative of each homologous pair. 4. During meiosis, each homologous chromosome pair is shuffled, so the resulting haploid cells each have a virtually unique combination of genes.

Meiosis typically consists of two nuclear and cytoplasmic divisions, designated the first and second meiotic divisions, or simply meiosis I and meiosis II (FIG. 10-15). Each includes prophase, metaphase, anaphase, and telophase stages. During meiosis I, the partner homologous chromosomes pair and then separate and move into different nuclei. In meiosis II, the sister chromatids that make up each duplicated chromosome separate from each other and are distributed to two different nuclei. The following discussion describes meiosis in an organism with a diploid chromosome number of 4. Refer to FIGURE 10-16 as you read.

Four haploid (n) cells, each with two unduplicated chromosomes

FIGURE 10-15 Overview of meiosis

MEIOSIS I Mid-prophase I

Late prophase I

Clare Hasenkampf / Biological Photo Service

INTERPHASE

Nucleolus Nuclear envelope Homologous chromosomes

Chromatin

Centrioles

Interphase preceding meiosis; DNA replicates.

MEIOSIS II Prophase II

Developing meiotic spindle

Homologous chromosomes synapse and exchange segments by crossing over; nuclear envelope breaks down.

Metaphase II

Anaphase II

Daughter chromosomes

Chromosomes condense again following brief period of interkinesis. DNA does not replicate again.

Chromosomes line up along cell's midplane.

Sister chromatids separate, and chromosomes move to opposite poles.

FIGURE 10-16 Animated Interphase and the stages of meiosis Meiosis consists of two nuclear divisions, meiosis I (top row) and meiosis II (bottom row). The LMs show sectioned plant cells, which lack centrioles. The drawings depict generalized animal cells with a diploid chromosome number of 4; the sizes of the nuclei and chromosomes are exaggerated to show the structures more clearly.

Metaphase I

Anaphase I

Telophase I

Cleavage furrow

Microtubule attached to kinetochore Separation of homologous chromosomes

Sister chromatids

Homologous chromosomes separate and move to opposite poles. Note that sister chromatids remain attached at their centromeres.

Telophase II

Four haploid cells

Nuclei form at opposite poles of each cell. Cytokinesis occurs.

Four gametes (animal) or four spores (plant) are produced.

25 μm

Tetrads line up on cell's midplane.Tetrads held together at chiasmata (sites of prior crossing-over).

FIGURE 10-16 Continued

One of each pair of homologous chromosomes is at each pole. Cytokinesis occurs.

D. Von Wettstein, Proceedings of the National Academy of Science 68: 851–855, 1971

Maternal sister chromatids

Chromosome

Paternal sister chromatids

Synaptonemal complex Chromosome Synaptonemal complex

0.5 μm

(b) TEM of a synaptonemal complex.

Chromatin Protein

Chromatin

Maternal sister chromatids

(a) A 3-D model of a tetrad with a complete synaptonemal complex.

FIGURE 10-17 A synaptonemal complex Synapsing homologous chromosomes in meiotic prophase I are held together by a synaptonemal complex, composed mainly of protein.

Prophase I includes synapsis and crossing-over As occurs during mitosis, the chromosomes duplicate in the S phase of interphase, before the complex movements of meiosis actually begin. Each duplicated chromosome consists of two chromatids, which are linked by cohesins. During prophase I, while the chromatids are still elongated and thin, the homologous chromosomes come to lie lengthwise side by side. This process is called synapsis, which means “fastening together.” For example, in an animal cell with a diploid number of four, synapsis results in two homologous pairs. One member of each homologous pair is called the maternal homologue because it was originally inherited from the female parent; the other member of a homologous pair is the paternal homologue because it was inherited from the male parent. Because each chromosome duplicated during interphase and now consists of two chromatids, synapsis results in the association of four chromatids. The resulting association is a tetrad. The number of tetrads per prophase I cell is equal to the haploid chromosome number. In an animal cell with a diploid number of 4, there are 2

tetrads (and a total of 8 chromatids); in a human cell at prophase I, there are 23 tetrads (and a total of 92 chromatids). Homologous chromosomes become closely associated during synapsis. Electron microscopic observations reveal that a characteristic structure, the synaptonemal complex, forms along the entire length of the synapsed homologues (FIG. 10-17). This proteinaceous structure holds the synapsed homologues together and is thought to play a role in chromosome crossing-over, a process in which enzymes break and rejoin DNA molecules, allowing paired homologous chromosomes to exchange genetic material. Crossing-over produces new combinations of genes. The genetic recombination from crossing-over greatly enhances the genetic variation—that is, new combinations of traits—among sexually produced offspring. Some biologists think that recombination is the main reason for sexual reproduction in eukaryotes. In addition to the unique processes of synapsis and crossingover, events similar to those in mitotic prophase also occur during prophase I. A spindle forms, consisting of microtubules and other components. In animal cells, one pair of centrioles moves to each pole, and astral microtubules form. The nuclear envelope disappears in late prophase I, and in cells with large and distinct chromosomes, the structure of the tetrads can be seen clearly with the microscope. The sister chromatids remain closely aligned along their lengths. However, the centromeres (and kinetochores) of the homologous chromosomes become separated from one another. In late prophase I, the homologous chromosomes are held together only at specific regions, termed chiasmata (sing., chiasma). Each chiasma originates at a crossing-over site, that is, a site at which homologous chromatids exchanged genetic material and rejoined, producing an X-shaped configuration (FIG. 10-18). At the chiasmata, cohesins hold homologous chromosomes together after the synaptonemal complex has been disassembled. Later, the cohesins dissociate from the chiasmata, freeing the homologous chromo-

Sister chromatids

Chiasmata

Sister chromatids

Chiasmata

Kinetochores Courtesy of J. Kezer

Kinetochores Sister chromatids 1 μm

(a) LM of a tetrad during late prophase I of a male meiotic cell (spermatocyte) from a salamander.

(b) A drawing showing the structure of the tetrad. The paternal chromatids are blue, and the maternal chromatids are red.

FIGURE 10-18 A meiotic tetrad with two chiasmata The two chiasmata are the result of separate crossing-over events.

some arms from one another. The consequences of crossing-over and genetic recombination are discussed in Chapter 11 (for example, see Fig. 11-12).

During meiosis I, homologous chromosomes separate Metaphase I occurs when the tetrads align on the midplane. Both sister kinetochores of one duplicated chromosome are attached by spindle fibers to the same pole, and both sister kinetochores of the other duplicated homologous chromosome are attached to the opposite pole. (By contrast, sister kinetochores of each duplicated chromosome are attached to opposite poles in mitosis.) During anaphase I, the paired homologous chromosomes separate, or disjoin, and move toward opposite poles. Each pole receives a random combination of maternal and paternal chromosomes, but only one member of each homologous pair is present at each pole. The sister chromatids remain united at their centromere regions. Again, this differs from mitotic anaphase, in which the sister chromatids separate and move to opposite poles. During telophase I, the chromatids generally decondense somewhat, the nuclear envelope may reorganize, and cytokinesis may take place. Each telophase I nucleus contains the haploid number of chromosomes, but each chromosome is a duplicated chromosome (it consists of a pair of chromatids). In our example, 2 duplicated chromosomes lie at each pole, for a total of 4 chromatids; humans have 23 duplicated chromosomes (46 chromatids) at each pole. An interphase-like stage called interkinesis usually follows. Interkinesis is not a true interphase: there is no S phase and therefore no DNA replication. Interkinesis is brief in most organisms and absent in some.

Chromatids separate in meiosis II Because the chromosomes usually remain partially condensed between divisions, the prophase of the second meiotic division is brief.

Prophase II is similar to mitotic prophase in many respects. There is no pairing of homologous chromosomes (indeed, only one member of each pair is present in each nucleus) and no crossing-over. During metaphase II the chromosomes line up on the midplanes of their cells. You can easily distinguish the first and second metaphases in diagrams; at metaphase I the chromatids are arranged in bundles of four (tetrads), and at metaphase II they are in groups of two (as in mitotic metaphase). This is not always so obvious in living cells. During anaphase II the chromatids, attached to spindle fibers at their kinetochores, separate and move to opposite poles, just as they would at mitotic anaphase. As in mitosis, each former chromatid is now referred to as a chromosome. Thus, at telophase II there is one representative for each homologous pair at each pole. Each is an unduplicated (single) chromosome. Nuclear envelopes re-form, the chromosomes gradually elongate to form chromatin fibers, and cytokinesis occurs. The two successive divisions of meiosis yield four haploid nuclei, each containing one of each kind of chromosome. Each resulting haploid cell has a different combination of genes. This genetic variation has two sources: (1) DNA segments are exchanged between maternal and paternal homologues during crossing-over. (2) During meiosis, the maternal and paternal chromosomes of homologous pairs separate independently. The chromosomes are “shuffled” so that each member of a pair becomes randomly distributed to one of the poles at anaphase I.

Mitosis and meiosis lead to contrasting outcomes Although mitosis and meiosis share many similar features, specific distinctions between these processes result in the formation of different types of cells. Mitosis is a single nuclear division in which sister chromatids separate from each other and are distributed to the two daughter cells, which are genetically identical to each other and to the original cell. A diploid cell that undergoes mitosis produces two diploid cells. Similarly, a haploid cell that

KEY POINT

Each species has a characteristic number of chromosomes that does not change. In each life cycle, the doubling of chromosomes that occurs during fertilization is compensated for by the reduction in chromosome number that occurs during meiosis.

Unicellular or multicellular haploid organism (n)

Gametes (n) Meiosis

Fertilization Mitosis

Mitosis

Zygote (2n) Gametes (n)

Mitosis Meiosis

Multicellular diploid organism (2n)

Fertilization Zygote (2n)

(a) Animals

(b) Most fungi and many protists

Gametophyte (n) (multicellular haploid organism) Mitosis

Mitosis

Spores (n)

Gametes (n)

Fertilization

Meiosis

Zygote (2n) Mitosis Sporophyte (2n) (multicellular diploid organism)

(c) Plants, some algae, and some fungi

FIGURE 10-19 Representative life cycles The color code and design here is used throughout the rest of the book. For example, in all life cycles the haploid (n) generation is shown in purple, and the diploid (2n) generation is gold. The processes of meiosis and fertilization always link the haploid and diploid generations.

undergoes mitosis produces two haploid cells. (Some eukaryotic organisms—certain yeasts, for example—are haploid, as are plants at certain stages of their life cycles.) Homologous chromosomes do not associate physically at any time in mitosis. In meiosis, a diploid cell undergoes two successive nuclear divisions, meiosis I and meiosis II. In prophase I of meiosis, the homologous chromosomes undergo synapsis to form tetrads. Homologous chromosomes separate during meiosis I, and sister chromatids separate during meiosis II. Meiosis ends with the formation of four genetically different, haploid daughter cells. The fates of these cells depend on the type of life cycle; in animals they differentiate as gametes, whereas in plants they become spores.

Review Are homologous chromosome pairs present in a diploid cell? Are they present in a haploid cell? How does the outcome of meiosis differ from the outcome of mitosis? Can haploid cells divide by mitosis? by meiosis?

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■ ■

10.5 SEXUAL LIFE CYCLES ■ ■ LEARNING OBJECTIVE 11 Compare the roles of mitosis and meiosis in various generalized life cycles.

Because sexual reproduction is characterized by the fusion of two haploid sex cells to form a diploid zygote, it follows that in a sexual life cycle, meiosis must occur before gametes can form. The timing of meiosis in the life cycle varies among species. In animals and a few other organisms, meiosis leads directly to gamete production (FIG. 10-19a on facing page). An organism’s somatic cells multiply by mitosis and are diploid; the only haploid cells produced are the gametes. Gametes develop when germ line cells, which give rise to the next generation, undergo meiosis. The formation of gametes is known as gametogenesis. Male gametogenesis, termed spermatogenesis, forms four haploid

■ ■

sperm cells for each cell that enters meiosis. (See Chapter 50 and Fig. 50-5 for a detailed description of spermatogenesis.) In contrast, female gametogenesis, called oogenesis, forms a single egg cell, or ovum, for every cell that enters meiosis. In this process, most of the cytoplasm goes to only one of the two cells produced during each meiotic division. At the end of meiosis I, one nucleus is retained and the other, called the first polar body, often degenerates. Similarly, at the end of meiosis II, one nucleus becomes another polar body and the other nucleus survives. In this way, one haploid nucleus receives most of the accumulated cytoplasm and nutrients from the original meiotic cell. (See Chapter 50 and Fig. 50-11 for a detailed description of oogenesis.) Although meiosis occurs at some point in a sexual life cycle, it does not always immediately precede gamete formation. Many eukaryotes, including most fungi and many protists, remain haploid (their cells dividing mitotically) throughout most of their life cycles, with individuals being unicellular or multicellular. Two haploid gametes (produced by mitosis) fuse to form a diploid zygote that undergoes meiosis to restore the haploid state (FIG. 10-19b). Examples of these types of life cycles are found in Figures 26-16 and 29-9. Plants and some algae and fungi have complicated life cycles (FIG. 10-19c). These life cycles, characterized by an alternation of generations, consist of a multicellular diploid stage, the sporophyte generation, and a multicellular haploid stage, the gametophyte generation. Sporophyte cells undergo meiosis to form haploid spores, each of which may divide mitotically to produce a multicellular haploid gametophyte. Gametophytes produce gametes by mitosis. The female and male gametes (egg and sperm cells) fuse to form a diploid zygote that divides mitotically to form a multicellular sporophyte. You can find more detailed descriptions of alternation of generations in plants in Chapters 27 and 28.

Review ■ ■

In animals, which cells are produced by mitosis? by meiosis? Assuming a sexually reproducing organism is haploid, how do meiosis and fertilization maintain the normal chromosome number?

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

10.1 (page 214) 1 Discuss the significance of chromosomes in terms of their information content. ■ Genes, the cell’s informational units, are made of DNA. In eukaryotes, DNA associates with protein to form the chromatin fibers that make up chromosomes.

2 Explain how DNA is packed into chromosomes in eukaryotic cells. ■

The organization of eukaryotic DNA into chromosomes allows the DNA to be accurately replicated and sorted into daughter cells without tangling. In eukaryotic cells DNA is associated with histones (basic proteins) to form nucleosomes, each of which consists of a histone bead with DNA wrapped around it. Nucleosomes are organized into large, coiled loops held together by nonhistone scaffolding proteins.

Watch cytokinesis unfold by clicking on the figure in CengageNOW.

10.2 (page 217) 3 Identify the stages in the eukaryotic cell cycle and describe their principal events. The eukaryotic cell cycle is the period from the beginning of one division to the beginning of the next. The cell cycle consists of interphase and M phase.

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10.3 (page 223) 6 Explain some ways in which the cell cycle is controlled. ■

TERPHASE IN

G1

S

Control mechanisms, called cell-cycle checkpoints, temporarily block key events from being initiated during the cell cycle. Cyclindependent kinases (Cdks) are protein kinases involved in regulating the cell cycle. Cdks are active only when they bind tightly to regulatory proteins called cyclins. Cyclin levels fluctuate predictably during the cell cycle.

10.4 (page 225) G2

7 Differentiate between asexual and sexual reproduction. Offspring produced by asexual reproduction usually have hereditary traits identical to those of the single parent. Mitosis is the basis for asexual reproduction in eukaryotic organisms. ■ In sexual reproduction, two haploid sex cells, or gametes, fuse to form a single diploid zygote. In a sexual life cycle, meiosis must occur before gametes can be produced. 8 Distinguish between haploid and diploid cells, and define homologous chromosomes. ■ A diploid cell has a characteristic number of chromosome pairs per cell. The members of each pair, called homologous chromosomes, are similar in length, shape, and other features and carry genes affecting the same kinds of attributes of the organism. ■ A haploid cell contains only one member of each homologous chromosome pair. 9 Explain the significance of meiosis and describe the process. ■ A diploid cell undergoing meiosis completes two successive cell divisions, yielding four haploid cells. Sexual life cycles in eukaryotes require meiosis, which makes it possible for each gamete to contain only half the number of chromosomes in the parent cell. ■ Meiosis I begins with prophase I, in which the members of a homologous pair of chromosomes physically join by the process of synapsis. Crossing-over is a process of genetic recombination during which homologous (nonsister) chromatids exchange segments of DNA strands. ■ At metaphase I, tetrads—each consisting of a pair of homologous chromosomes held together by one or more chiasmata—line up on the metaphase plate. The members of each pair of homologous chromosomes separate during meiotic anaphase I and are distributed to different nuclei. Each nucleus contains the haploid number of chromosomes; each chromosome consists of two chromatids. ■ During meiosis II, the two chromatids of each chromosome separate, and one is distributed to each daughter cell. Each former chromatid is now a chromosome. ■

M PHASE ■

■

Interphase consists of the first gap phase (G1), the synthesis phase (S), and the second gap phase (G2). During the G1 phase, the cell grows and prepares for the S phase. During the S phase, DNA and the chromosome proteins are synthesized, and chromosome duplication occurs. During the G2 phase, protein synthesis increases in preparation for cell division. M phase consists of mitosis, the nuclear division that produces two nuclei identical to the parental nucleus, and cytokinesis, the division of the cytoplasm to yield two daughter cells. Learn more about the cell cycle by clicking on the figure in CengageNOW.

4 Describe the structure of a duplicated chromosome, including the sister chromatids, centromeres, and kinetochores. A duplicated chromosome consists of a pair of sister chromatids, which contain identical DNA sequences. Each chromatid includes a constricted region called a centromere. Sister chromatids are tightly associated in the region of their centromeres. Attached to each centromere is a kinetochore, a protein structure to which microtubules can bind. 5 Explain the significance of mitosis and describe the process. ■ Mitosis assures that the chromosome number is preserved when one eukaryotic cell divides to form two. In mitosis, identical chromosomes are distributed to each pole of the cell, and a nuclear envelope forms around each set. ■ During prophase, the structure of the duplicated chromosomes becomes apparent as the chromatin condenses; each is composed of a pair of identical sister chromatids. The nuclear envelope begins to disassemble, and the mitotic spindle begins to form. ■ During prometaphase, spindle microtubules attach to kinetochores of chromosomes, and chromosomes begin to move toward the cell’s midplane. ■ During metaphase, the chromosomes are aligned on the cell’s midplane, or metaphase plate; the mitotic spindle is complete, and the kinetochores of the sister chromatids are attached by microtubules to opposite poles of the cell. ■ During anaphase, the sister chromatids separate and move to opposite poles. Each former chromatid is now a chromosome. ■ During telophase, a nuclear envelope re-forms around each set of chromosomes, nucleoli become apparent, the chromosomes uncoil, and the spindle disappears. Cytokinesis generally begins in telophase. ■

Walk step-by-step through the stages of mitosis by clicking on the figure in CengageNOW.

Walk step-by-step through the stages of meiosis by clicking on the figure in CengageNOW.

10 Contrast mitosis and meiosis, emphasizing the different outcomes. ■

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Mitosis involves a single nuclear division in which the two daughter cells formed are genetically identical to each other and to the original cell. Synapsis of homologous chromosomes does not occur during mitosis. Meiosis involves two successive nuclear divisions and forms four haploid cells. Synapsis of homologous chromosomes occurs during prophase I of meiosis. Watch a movie that features living cells undergoing mitosis and meiosis in CengageNOW.

10.5 (page 233) 11 Compare the roles of mitosis and meiosis in various generalized life cycles. Somatic cells of animals are diploid and are produced by mitosis. The only haploid cells are gametes, produced by gametogenesis, which in animals occurs by meiosis. ■ Most fungi and many protists are haploid and are produced by mitosis. The only diploid stage is the zygote, which undergoes meiosis to restore the haploid state. ■

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The life cycle of plants, some algae, and some fungi includes an alternation of generations. The multicellular diploid sporophyte generation forms haploid spores by meiosis. Each spore divides mitotically to form a multicellular haploid gametophyte generation, which produces gametes by mitosis. Two haploid gametes then fuse to form a diploid zygote, which divides mitotically to produce a new sporophyte generation.

T E S T YO U R U N D E R S TA N D I N G 1. Chromatin fibers include (a) DNA and structural polysaccharides (b) RNA and phospholipids (c) protein and carbohydrate (d) DNA and protein (e) triacylglycerol and steroids 2. A nucleosome consists of (a) DNA and scaffolding proteins (b) scaffolding proteins and histones (c) DNA and histones (d) DNA, histones, and scaffolding proteins (e) histones only 3. The term S phase refers to (a) DNA synthesis during interphase (b) synthesis of chromosome proteins during prophase (c) gametogenesis in animal cells (d) synapsis of homologous chromosomes (e) fusion of gametes in sexual reproduction

11. A chiasma links a pair of (a) homologous chromosomes at prophase II (b) homologous chromosomes at late prophase I (c) sister chromatids at metaphase II (d) sister chromatids at mitotic metaphase (e) sister chromatids at metaphase I 12. Match the following descriptions (a–e) to the steps of meiosis (1–5) in the following figure. (a) haploid cells with unduplicated chromosomes (b) haploid cells with duplicated chromosomes (c) synapsis of homologous chromosomes to form tetrads (d) sister chromatids move to opposite poles (e) homologous chromosomes move to opposite poles

4. At which of the following stages do human skin cell nuclei have the same DNA content? (a) early mitotic prophase and late mitotic telophase (b) G1 and G2 (c) G1 and early mitotic prophase (d) G1 and late mitotic telophase (e) G2 and late mitotic telophase

1

5. In a cell at _____, each chromosome consists of a pair of attached chromatids. (a) mitotic prophase (b) meiotic prophase II (c) meiotic prophase I (d) meiotic anaphase I (e) all of the preceding

2

6. The molecular tether that links sister chromatids of a duplicated chromosome to each other is (a) condensin (b) actin (c) myosin (d) cohesin (e) actomyosin 3

7. In an animal cell at mitotic metaphase, you would expect to find (a) two pairs of centrioles located on the metaphase plate (b) a pair of centrioles inside the nucleus (c) a pair of centrioles within each microtubule-organizing center (d) a centriole within each centromere (e) no centrioles

4

8. Cell plate formation usually begins during (a) telophase in a plant cell (b) telophase in an animal cell (c) G2 in a plant cell (d) G2 in an animal cell (e) a and b 9. A diploid nucleus at early mitotic prophase has _____ set(s) of chromosomes; a diploid nucleus at mitotic telophase has _____ set(s) of chromosomes. (a) 1; 1 (b) 1; 2 (c) 2; 2 (d) 2; 1 (e) not enough information has been given 10. You would expect to find a synaptonemal complex in a cell at (a) mitotic prophase (b) meiotic prophase I (c) meiotic prophase II (d) meiotic anaphase I (e) meiotic anaphase II

5

CRITICAL THINKING 1. How can two species have the same chromosome number yet have very different attributes? 2. How does the DNA content of the cell change from the beginning of interphase to the end of interphase? Does the number of chromatids change? Explain. Does the number of chromosomes change? Explain. In questions 3–4, decide whether each is an example of sexual or asexual reproduction, and state why. 3. A diploid queen honeybee produces haploid eggs by meiosis. Some of these eggs are never fertilized and develop into haploid male honeybees (drones). 4. Seeds develop after a flower has been pollinated with pollen from the same plant. 5. ANALYZING DATA. In the experiment with spindle microtubules during anaphase (see Fig. 10-10), how would the spindle mi-

crotubules have looked during anaphase if they had disassembled at their polar ends instead of their kinetochore ends? 6. EVOLUTION LINK. How does mitosis provide evidence for relationships among eukaryotes as diverse as mammals and seaweeds? 7. EVOLUTION LINK. Some organisms—for example, certain fungi—reproduce asexually when the environment is favorable and sexually when the environment becomes unfavorable. What might be the evolutionary advantage of sexual reproduction with the associated process of meiosis during unfavorable conditions? Additional questions are available in CengageNOW at www.cengage.com/ login.

11

The Basic Principles of Heredity

© Science Source/Photo Researchers, Inc.

Gregor Mendel. This painting shows Mendel with his pea plants in the monastery garden at Brünn, Austria (now Brüno, Czech Republic).

KEY CONCEPTS

H

uman characters such as eye color and hair color, along with a multitude of other characteristics, are passed on from one generation

11.1 The experiments of Gregor Mendel, a pioneer in the

to another. Heredity, the transmission of genetic information from par-

field of genetics, revealed the basic principles of inheritance. In Mendel’s principle of segregation, members of a gene pair segregate (separate) from one another prior to gamete formation. In the principle of independent assortment, members of different gene pairs assort independently (randomly) into gametes.

ent to offspring, generally follows predictable patterns in organisms as

11.2 You can use probability to predict Mendelian inheritance: the product rule shows how to combine the probabilities of independent events, and the sum rule shows how to combine the probabilities of mutually exclusive events. 11.3 Chromosome behavior during meiosis helps explain Mendel’s principles of inheritance.

11.4 Distinctive inheritance patterns (that is, “extensions” to Mendel’s principles) characterize some traits.

diverse as humans, penguins, baker’s yeast, and sunflowers. Genetics, the science of heredity, studies both genetic similarities and genetic variation, the differences between parents and offspring or among individuals of a population. The study of inheritance as a modern branch of science began in the mid-19th century with the work of Gregor Mendel (1822–1884), a monk who bred pea plants (see picture). Mendel was the first scientist to effectively apply quantitative methods to the study of inheritance. He did not merely describe his observations; he planned his experiments carefully,

recorded the data, and analyzed the results mathematically. Although unappreciated during his lifetime, his work was rediscovered in 1900. During the decades following the rediscovery of Mendel’s find-

11.1 MENDEL’S PRINCIPLES OF INHERITANCE

ings, geneticists extended Mendel’s principles by correlating the

■ ■ LEARNING OBJECTIVES

transmission of genetic information from generation to generation

1

with the behavior of chromosomes during meiosis. By studying a variety of organisms, geneticists verified Mendel’s findings and added a growing list of so-called exceptions to his principles. Geneticists study not only the transmission of genes but also

2 3 4

the expression of genetic information. As you will see in this chapter and those that follow, understanding the relationships between an organism’s genes and its characteristics has become increasingly sophisticated as biologists have learned more about the orderly transmission of information in cells from one generation to another.

Define the terms phenotype, genotype, locus, allele, dominant allele, recessive allele, homozygous, and heterozygous. Describe Mendel’s principles of segregation and independent assortment. Distinguish among monohybrid, dihybrid, and test crosses. Explain Mendel’s principles of segregation and independent assortment, given what scientists now know about genes and chromosomes.

Gregor Mendel was not the first plant breeder. At the time he began his work, breeders had long recognized the existence of hybrid plants and animals, the offspring of two genetically dissimilar parents. When Mendel began his breeding experiments in 1856, two

RESEARCH METHOD Why Is It Used?

How Is It Done?

Garden peas normally self-fertilize during reproduction, that is, the male and female gametes are from the same flower. Because pea petals completely enclose the reproductive parts, there is little chance of natural cross-pollination between separate flowers. Cross-pollination enables the researcher to study various patterns of inheritance in peas. 1

Reproductive structures of flowers enclosed by petals

3

Anthers snipped from the flower

4

Pollen from a different flower brushed onto tip of carpel

5

Fertilized carpel produces seeds, which are planted

Anther Carpel

Carpel 2

6

Petals opened to reveal male and female reproductive structures

Offspring observed

FIGURE 11-1 How garden peas are cross-pollinated

main concepts about inheritance were widely accepted: (1) All hybrid plants that are the offspring of genetically pure, or truebreeding, parents are similar in appearance. (2) When these hybrids mate with each other, they do not breed true; their offspring show a mixture of traits. Some look like their parents, and some have features like those of their grandparents. Mendel’s genius lay in his ability to recognize a pattern in the way the parental traits reappear in the offspring of hybrids. Before Mendel, no one had categorized and counted the offspring and analyzed these regular patterns over several generations to the extent he did. Just as geneticists do today, Mendel chose the organism for his experiments very carefully. The garden pea, Pisum sativum, had several advantages. Pea plants are easy to grow, and many varieties were commercially available. Another advantage of pea plants is that controlled pollinations are relatively easy to conduct. Pea flowers have both male and female parts and naturally self-pollinate. However, the anthers (the male parts of the flower that produce pollen) can be removed to prevent self-fertilization (FIG. 11-1). Pollen from a different source can then be applied to the stigma (the receptive surface of the carpel, or female part). Pea flowers are easily protected from other sources of pollen because the petals completely enclose the reproductive structures. Mendel obtained his original pea seeds from commercial sources and did some important preliminary work before starting

Flower color

Seed color

Yellow

his actual experiments. For two years he verified that the varieties were true-breeding lines for various inherited features. Today, scientists use the term phenotype to refer to the physical appearance of an organism and genotype to refer to the genetic makeup for that organism, which is most often expressed in symbols. A true-breeding line produces only offspring expressing the same phenotype (for example, round seeds or tall plants) generation after generation. During this time Mendel apparently chose those traits of his pea strains that he could study most easily. He probably made the initial observations that later formed the basis of his hypotheses. Mendel eventually chose strains representing seven characters, the attributes (such as seed color) for which heritable differences, or traits, are known (such as yellow seeds and green seeds). The characters Mendel selected had clearly contrasting phenotypes (FIG. 11-2). Mendel’s results were easy to analyze because he chose easily distinguishable phenotypes and limited the genetic variation studied in each experiment. Mendel began his experiments by crossing plants from two different true-breeding lines with contrasting phenotypes; these genetically pure individuals constituted the parental generation, or P generation. In every case, the members of the first generation of offspring all looked alike and resembled one of the two parents. For example, when he crossed tall plants with short plants, all the

Pod color

Pod shape

Green

Purple Seed shape

Smooth Wrinkled White

Green

Yellow

Inflated

Stem height

Tall

Short

FIGURE 11-2 Seven characters in Mendel’s study of pea plants Each character had two clearly distinguishable phenotypes; the dominant phenotype is boldface.

Pinched

Flower position

Axial

Terminal

KEY EXPERIMENT QUESTION: When the F1 generation of tall pea plants is selfpollinated, what phenotypes appear in the F2 generation? HYPOTHESIS: Although only the “factor” (gene) for tall height is expressed in the F1 generation, Mendel hypothesized that the factor for short height is not lost. He predicted that the short phenotype would reappear in the F2 generation. EXPERIMENT: Mendel crossed true-breeding tall pea plants with true-breeding short pea plants, yielding only tall offspring in the F1 generation. He then allowed these F1 individuals to self-pollinate to yield the F2 generation. P Generation

X

Tall plant T

Short plant

T

t

t

F1 Generation

All tall plants T

t

F2 Generation

offspring were tall (FIG. 11-3). These offspring were the first filial generation, or the F1 generation (filial is from the Latin for “sons and daughters”). The second filial generation, or F2 generation, resulted from a cross between F1 individuals or by self-pollination of F1 individuals. Mendel’s F2 generation in this experiment included 787 tall plants and 277 short plants. TABLE 11-1 shows Mendel’s experimental results for all seven pea characters. Most breeders in Mendel’s time thought that inheritance involved the blending of traits. In blending inheritance, male and female gametes supposedly contained fluids that blended together during reproduction to produce hybrid offspring with features intermediate between those of the mother and father. In fact, some plant breeders had obtained such hybrids. Although Mendel observed some intermediate types of hybrids, he chose for further study those F1 hybrids in which “hereditary factors” (as he called them) from one of the parents apparently masked the expression of those factors from the other parent. Other breeders had also observed these types of hybrids, but they had not explained them. Using modern terms, we say that the factor expressed in the F1 generation (tallness, in our example) is dominant; the one hidden in the F1 (shortness) is recessive. Dominant traits mask recessive ones when both are present in the same individual. Although scientists know today that dominance is not always observed, the fact that dominance can occur was not consistent with the notion of blending inheritance. Mendel’s results were inconsistent with the hypothesis of blending inheritance in a more compelling way. Once two fluids have blended, it is very difficult to imagine how they can separate. However, in the preceding example, in the F1 generation the hereditary factor(s) that controlled shortness clearly was not lost or blended inseparably with the hereditary factor(s) that controlled tallness because shortness reappeared in the F2 generation. Mendel was very comfortable with the theoretical side of biology because he was also a student of physics and mathematics. He therefore proposed that each kind of inherited feature of an organism is controlled by two factors that behave like discrete particles and are present in every individual. To Mendel these hereditary factors were abstractions—he knew nothing about chromosomes and DNA. These factors are essentially what scientists today call genes—units of heredity that affect an organism’s traits. At the molecular level, a gene is a DNA sequence that contains the information to make an RNA or pro-

TABLE 11-1

Mendel’s Experimental Results for Seven Characters

Characters and Traits in Peas Tall plant T

T

Tall plant T

Tall plant

t

T

t

Short plant t

t

3 tall : 1 short

RESULTS AND CONCLUSION: The F2 generation included 787 tall and 277 short plants, which resulted in a ratio of about 3:1. Thus, Mendelian traits pass to successive generations in fixed ratios.

FIGURE 11-3 Animated One of Mendel’s pea crosses

Character

Dominant Trait ×

Stem height Flower color Flower position Pod color Pod shape Seed color Seed shape

Tall Purple Axial Green Inflated Yellow Smooth

× × × × × × ×

F2 Generation

Recessive Trait

Dominant: Recessive

Ratio

Short White Terminal Yellow Pinched Green Wrinkled

787:277 705:224 651:207 428:152 882:299 6022:2001 5474:1850

2.84:1 3.15:1 3.14:1 2.82:1 2.95:1 3.01:1 2.96:1

tein product with a specific function. (Chapter 13 traces the development of ideas regarding the nature of genes.) Mendel’s experiments led to his discovery and explanation of the major principles of heredity, which we now know as the principles of segregation and independent assortment (see TABLE 11-2 for a summary of Mendel’s model of inheritance). We discuss the first principle next and the second later in the chapter.

Alleles separate before gametes are formed: the principle of segregation Alternative forms of a gene are called alleles. In the example in Figure 11-3, each F1 generation tall plant had two different alleles that control plant height: a dominant allele for tallness (which we designate T) and a recessive allele for shortness (designated t). Because the tall allele was dominant, these plants were tall. To explain his experimental results, Mendel proposed an idea now known as the principle of segregation. Using modern terminology, the principle of segregation states that before sexual reproduction occurs, the two alleles carried by an individual parent must become separated (that is, segregated). Recall that during meiosis, homologous chromosomes, and therefore the alleles that reside on them, separate. (You may want to review meiosis in Figure 10-16.) As a result, each sex cell (egg or sperm) formed contains only one allele of each pair. (Later, at the time of fertilization, each haploid gamete contributes one chromosome from each homologous pair and therefore one allele for each gene pair.) An essential feature of meiosis is that the alleles remain

TABLE 11-2

intact (one does not mix with or eliminate the other); thus, recessive alleles are not lost and can reappear in the F2 generation. In our example, before the F1 plants formed gametes, the allele for tallness segregated from the allele for shortness; so half the gametes contained a T allele, and the other half, a t allele (FIG. 11-4). The random process of fertilization led to three possible combinations of alleles in the F2 offspring: one-fourth with two tallness alleles (TT), one-fourth with two shortness alleles (tt), and one-half with one allele for tallness and one for shortness (Tt). Because both TT and Tt plants are tall, Mendel expected approximately three-fourths (787 of the 1064 plants he obtained) to express the phenotype of the dominant allele (tall) and about one-fourth (277/1064) to express the phenotype of the recessive allele (short). We will explain the mathematical reasoning behind these predictions later in the chapter.

KEY POINT

Mendel’s principle of segregation is related to the events of meiosis: the separation of homologous chromosomes during meiosis results in the segregation of alleles.

Homologous chromosomes METAPHASE I

T

Mendel’s Model of Inheritance

T t

t

1. Alternative forms of a “factor” (what we now call a gene) account for variations in inherited traits. Although Mendel observed only two forms (what we now call alleles) for each factor he studied, we now know that many genes have more than two alleles.

METAPHASE II

2. Inherited traits pass from parents to offspring as unmodified factors. Mendel did not observe offspring of intermediate appearance, as a hypothesis of blending inheritance would have predicted. Exceptions to this concept are known today.

T

t

T

t

3. Each individual has two sets of factors, one of each pair inherited from the mother and one from the father. It does not matter which parent contributes which set of factors. 4. The paired factors separate during the formation of reproductive cells (the principle of segregation). Because of meiosis, which was discovered after Mendel’s time, each parent passes one set of factors to each offspring. 5. Factors may be expressed or hidden in a given generation, but they are never lost. For example, factors not expressed in the F1 generation reappear in some F2 individuals. 6. Each factor is passed to the next generation independently from all other factors (the principle of independent assortment). Research since Mendel’s time has revealed that there are exceptions to this principle.

T

T

T-carrying gametes

t

t

t-carrying gametes

FIGURE 11-4 Animated Chromosomes and segregation Note that half of the gametes carry T and half carry t.

Alleles occupy corresponding loci on homologous chromosomes Today scientists know that each unduplicated chromosome consists of one long, linear DNA molecule and that each gene is actually a segment of that DNA molecule. We also know that homologous chromosomes not only are similar in size and shape but also usually have the same genes (often with different alleles) located in corresponding positions. The term locus (pl., loci) originally designated the location of a particular gene on the chromosome (FIG. 11-5). We are actually referring to a segment of the DNA that has the information for controlling some aspect of the structure or function of the organism. One locus may govern seed color, another seed shape, another shape of the pods, and so on. Traditional genetic methods can infer the existence of a particular locus only if at least two allelic variants of that locus, producing contrasting phenotypes (for

A gamete has one set of chromosomes, the n number. It carries one chromosome of each homologous pair. A given gamete can have only one gene of any particular pair of alleles.

When the gametes fuse, the resulting zygote is diploid (2n) and has homologous pairs of chromosomes. For purposes of illustration, these are shown physically paired.

(a) One member of each pair of homologous chromosomes is of maternal origin (red), and the other is paternal (blue).

example, yellow peas versus green peas), are available for study. In the simplest cases, an individual can express one (yellow) or the other (green) but not both. Alleles are, therefore, genes that govern variations of the same character (yellow versus green seed color) and occupy corresponding loci on homologous chromosomes. Geneticists assign each allele of a locus a single letter or group of letters as its symbol.1 Although geneticists often use more complicated forms of notation, it is customary when working simple genetics problems to indicate a dominant allele with a capital letter and a recessive allele with the same letter in lowercase. Remember that the term locus designates not only a position on a chromosome but also a type of gene controlling a particular character; thus, Y (yellow) and y (green) represent a specific pair of alleles of a locus involved in determining seed color in peas. Although you may initially be uncomfortable with the fact that geneticists sometimes use the term gene to specify a locus and at other times to specify one of the alleles of that locus, the meaning is usually clear from the context.

A monohybrid cross involves individuals with different alleles of a given locus The basic principles of genetics and the use of genetics terms are best illustrated by examples. In the simplest case, a monohybrid cross, the inheritance of two different alleles of a single locus, is studied. FIGURE 11-6 illustrates a monohybrid cross featuring a locus that governs coat color in guinea pigs. The female comes from 1 Early geneticists developed their own symbols to represent genes and alleles. Later, groups of scientists met and decided on specific symbols for a given research organism, such as the fruit fly, but each research group had its own rules for assigning symbols. Universally accepted rules for assigning symbols for genes and alleles still do not exist.

Gene loci A pair of alleles

Alleles controlling fur color: Black Brown

Alleles controlling fur length: Long Short These genes are not allelic to one another

(b) These chromosomes are nonhomologous. Each chromosome is made up of hundreds or thousands of genes. A locus is the specific place on a chromosome where a gene is located.

(c) These chromosomes are homologous. Alleles are members of a gene pair that occupy corresponding loci on homologous chromosomes.

FIGURE 11-5 Animated Gene loci and their alleles

(d) Alleles govern the same character but do not necessarily contain the same information.

a true-breeding line of black guinea pigs. We say she is homozygous for black because the two alleles she carries for this locus are identical. The brown male is also from a true-breeding line and is homozygous for brown. What color would you expect the F1 offspring to be? Actually, it is impossible to make such a prediction without more information. In this particular case, the F1 offspring are black, but they are heterozygous, meaning they carry two different alleles for this locus. The brown allele determines coat color only in a homozygous brown individual; it is a recessive allele. The black allele influences coat color in both homozygous black and heterozygous individuals; it is a dominant allele. On the basis of this information, we can use symbols to designate the dominant black allele B and the recessive brown allele b. During meiosis in the female parent (BB), the two B alleles separate according to Mendel’s principle of segregation, so each egg has only one B allele. In the male (bb) the two b alleles sepa-

Black female (BB)

Brown male (bb)

P generation

Gametes b

B

All Bb Dominant B masks recessive b

F1 generation

Gametes from F1 female 1 2 Gametes from F1 male 1 2

1 2

B

1 2

b

1 BB 4

1 Bb 4

1 Bb 4

1 bb 4

B

b

F2 generation

FIGURE 11-6 Animated A monohybrid cross in guinea pigs In this example, a homozygous black guinea pig is mated with a homozygous brown guinea pig. The F1 generation includes only black individuals. However, the mating of two of these F1 offspring yields F2 generation offspring in the expected ratio of 3 black to 1 brown, indicating that the F1 individuals are heterozygous.

rate, so each sperm has only one b allele. The fertilization of each B egg by a b sperm results in heterozygous F1 offspring, each Bb; that is, each individual has one allele for brown coat and one for black coat. Because this is the only possible combination of alleles present in the eggs and sperm, all the F1 offspring are Bb. Also, note that it does not make any difference whether the offspring receive the dominant allele from the mother or father guinea pig. A Punnett square predicts the ratios of the various offspring of a cross During meiosis in heterozygous black guinea pigs (Bb), the chromosome containing the B allele becomes separated from its homologue (the chromosome containing the b allele), so each normal sperm or egg contains B or b but never both. Heterozygous Bb individuals form gametes containing B alleles and gametes containing b alleles in equal numbers. Because no special attraction or repulsion occurs between an egg and a sperm containing the same allele, fertilization is a random process. As you can see in Figure 11-6, the possible combinations of eggs and sperm at fertilization can be represented in the form of a grid, or Punnett square, devised by the early English geneticist Sir Reginald Punnett. The types of gametes (and their expected frequencies) from one parent are listed across the top, and those from the other parent are listed along the left side. The squares are then filled in with the resulting F2 combinations. Three-fourths of all F2 offspring have the genetic constitution BB or Bb and are phenotypically black; one-fourth have the genetic constitution bb and are phenotypically brown. The genetic mechanism that governs the approximate 3:1 F2 ratios obtained by Mendel in his pea-breeding experiments is again evident. These ratios are called monohybrid F2 phenotypic ratios. The phenotype of an individual does not always reveal its genotype As mentioned earlier, an organism’s phenotype is its appearance with respect to a certain inherited trait. However, because some alleles may be dominant and others recessive, we cannot always determine, simply by examining its phenotype, which alleles are carried by an organism. In the cross we have been considering, the genotype of the female parent is homozygous dominant, BB, and her phenotype is black. The genotype of the male parent is homozygous recessive, bb, and his phenotype is brown. The genotype of all the F1 offspring is heterozygous, Bb, and their phenotype is black. To prevent confusion, we always indicate the genotype of a heterozygous individual by writing the symbol for the dominant allele first and the recessive allele second (always Bb, never bB). The expression of dominance partly explains why an individual may resemble one parent more than the other, even though the two parents contribute equally to their offspring’s genetic constitution. Dominance is not predictable—that is, dominance is not an intrinsic part of an allele but is the property of an allele relative to other alleles. Dominance is produced by the mechanism of gene expression and can be determined only by experiment. In one species of animal, black coat may be dominant to brown; in another species, brown may be dominant to black. In a population, the dominant phenotype is not necessarily more common than the recessive phenotype.

KEY EXPERIMENT QUESTION: How do you determine the genotype of an individual with a dominant phenotype? HYPOTHESIS: A test cross between an individual with a dominant phenotype but unknown genotype and an individual that is homozygous recessive for the gene in question yields offspring that provide insight into the genotype of the dominant individual. EXPERIMENT: Cross a black guinea pig of unknown genotype with a brown guinea pig, which is known to be homozygous recessive.

?

If homozygous Black (BB)

?

Homozygous Brown (bb)

If heterozygous Black (Bb)

Gametes

Gametes B

B

b

All Bb

Homozygous Brown (bb)

b

b

Eggs

All offspring are black and heterozygous

b Sperm

(a) If a black guinea pig is mated with a brown guinea pig and all the offspring are black, the black parent probably has a homozygous genotype.

1 2

1 2

1 2

Bb Heterozygous black

1 2

bb Homozygous brown

B

b

(b) If any of the offspring is brown, the black guinea pig must be heterozygous. The expected phenotypic ratio is 1 black to 1 brown. RESULTS AND CONCLUSION: There are two possible outcomes. If all the offspring are black, the black parent probably has a homozygous genotype. If any of the offspring are brown, the black parent must be heterozygous. The expected ratio is 1 black: 1 brown.

FIGURE 11-7 Animated A test cross in guinea pigs

A test cross can detect heterozygosity Guinea pigs with the genotypes BB and Bb are alike phenotypically; they both have black coats. How, then, can we know the genotype of a black guinea pig? Geneticists accomplish this by performing an experimental cross known as a test cross, in which an individual of unknown genotype is crossed with a recessive individual (FIG. 11-7). In a test cross, the alleles carried by the gametes from the parent of unknown genotype are never “hidden” in the offspring by dominant alleles contributed by the other parent. Therefore, you can deduce the genotypes of all offspring directly

from their phenotypes. If all the offspring were black, what inference would you make about the genotype of the black parent? If any of the offspring were brown, what conclusion would you draw regarding the genotype of the black parent? Would you be more certain about one of these inferences than about the other?2 2

If any of the offspring were brown, the black parent must be heterozygous, Bb. If all the offspring were black, you could infer that the black parent is probably homozygous, BB. (For example, statisticians calculate that if the black parent is 1 heterozygous, the probability of four black offspring, and no brown, is only — 16 .)

Mendel conducted several test crosses; for example, he bred F1 (tall) pea plants with homozygous recessive (tt) short ones. He reasoned that the F1 individuals were heterozygous (Tt) and would be expected to produce equal numbers of T and t gametes. Because the homozygous short parents (tt) were expected to produce only t gametes, Mendel hypothesized that he would obtain equal numbers of tall (Tt) and short (tt) offspring. His results agreed with his hypothesis, providing additional evidence for the hypothesis that there is 1:1 segregation of the alleles of a heterozygous parent. Thus, Mendel’s principle of segregation not only explained the known facts, such as the 3:1 monohybrid F2 phenotypic ratio, but also let him successfully anticipate the results of other experiments—in this case, the 1:1 test-cross phenotypic ratio.

A dihybrid cross involves individuals that have different alleles at two loci Monohybrid crosses involve a pair of alleles of a single locus. Mendel also analyzed crosses involving alleles of two or more loci. A mating between individuals with different alleles at two loci is called a dihybrid cross. Consider the case of two pairs of alleles carried on nonhomologous chromosomes (that is, one pair of alleles is in one pair of homologous chromosomes, and the other pair of alleles is in a different pair of homologous chromosomes). Each pair of alleles is inherited independently; that is, each pair segregates during meiosis independently of the other. An example of a dihybrid cross carried through the F2 generation is shown in FIGURE 11-8. In this example, black is dominant to brown, and short hair is dominant to long hair. When a homozygous, black, short-haired guinea pig (BBSS) and a homozygous, brown, long-haired guinea pig (bbss) are mated, the BBSS animal produces gametes that are all BS, and the bbss individual produces gametes that are all bs. Each gamete contains one allele for each of the two loci. The union of the BS and bs gametes yields only individuals with the genotype BbSs. All these F1 offspring are heterozygous for hair color and for hair length, and all are phenotypically black and short-haired. Each F1 guinea pig produces four kinds of gametes with equal probability: BS, Bs, bS, and bs. Hence, the Punnett square has 16 (that is, 42) squares representing the F2 offspring, some of which are genotypically or phenotypically alike. There are 9 chances in 16 of obtaining a black, short-haired individual; 3 chances in 16 of obtaining a black, long-haired individual; 3 chances in 16 of obtaining a brown, short-haired individual; and 1 chance in 16 of obtaining a brown, long-haired individual. This 9:3:3:1 phenotypic ratio is expected in a dihybrid F2 if the hair color and hair length loci are on nonhomologous chromosomes.

Black, short-haired

Brown, long-haired

BBSS

bbss

P generation

Gametes BS

bs

F1 generation All BbSs Gametes formed by segregation and independent assortment of alleles Gametes from F1 female 1 4

Gametes from F1 male 1 4

1 4 1 4 1 4

BS

1 4

1 4

Bs

bS

1 4

bs

BBSS

BBSs

BbSS

BbSs

BS

Black, short

Black, short

Black, short

Black, short

BBSs

BBss

BbSs

Bbss

Bs

Black, short

Black, long

Black, short

Black, long

BbSS

BbSs

bbSS

bbSs

bS

Black, short

Black, short

Brown, short

Brown, short

BbSs

Bbss

bbSs

bbss

bs

Black, short

Black, long

Brown, short

Brown, long

F2 generation F2 phenotypes 9 16 Black, short-haired

3 16

3 16

1 16

Black, long-haired

Brown, short-haired

Brown, long-haired

FIGURE 11-8 Animated A dihybrid cross in guinea pigs

Alleles on nonhomologous chromosomes are randomly distributed into gametes: the principle of independent assortment

When a black, short-haired guinea pig is crossed with a brown, long-haired one, all the offspring are black and have short hair. However, when two members of the F1 generation are crossed, the ratio of phenotypes is 9:3:3:1.

On the basis of results similar to the guinea pig example, Mendel formulated the principle of inheritance, now known as Mendel’s principle of independent assortment, which states that members of any gene pair segregate from one another independently of the

members of the other gene pairs. This mechanism occurs in a regular way to ensure that each gamete contains one allele for each locus, but the alleles of different loci are assorted at random with respect

to each other in the gametes. The independent assortment of these alleles can result in genetic recombination (or simply recombination), the process of assorting and passing alleles to offspring in new combinations that are different from those in the parents. Today we recognize that independent assortment is related to the events of meiosis. It occurs because two pairs of homologous chromosomes can be arranged in two different ways at metaphase I of meiosis (FIG. 11-9). These arrangements occur randomly, with approximately half the meiotic cells oriented one way and half oriented the opposite way. The orientation of the homologous chromosomes on the metaphase plate then determines the way they subsequently separate and disperse into the haploid cells. (As you will soon see, however, independent assortment does not always occur.)

KEY POINT

Recognition of Mendel’s work came during the early 20th century Mendel reported his findings at a meeting of the Brünn Society for the Study of Natural Science (in what is now the Czech Republic); he published his results in the society’s report in 1866. At that time biology was largely a descriptive science, and biologists had little interest in applying quantitative and experimental methods such as those Mendel had used. Other biologists of the time did not appreciate the importance of his results nor his interpretations of those results. For 34 years his findings were largely neglected. In 1900, Hugo DeVries in Holland, Carl Correns in Germany, and Erich von Tschermak in Austria recognized Mendel’s princi-

Mendel’s principle of independent assortment—that factors for different characteristics separate independently of one another prior to gamete formation—is a direct consequence of the events of meiosis.

b

bB

B

b

bB

B

S

Ss

s

METAPHASE I

s

s S

S

METAPHASE II

b

b

b

B

B

b

b

B

B

s

s

S

S

S

S

s

s

b

s

bs

s

B

B

S

S

BS

b

b

S

bS

S

B

B

s

s

Bs

FIGURE 11-9 Animated Meiosis and independent assortment Two different pairs of homologous chromosomes can line up two different ways at metaphase I and be subsequently distributed. A cell with the orientation shown at the left produces half BS and half bs gametes.

Conversely, the cell at the right produces half Bs and half bS gametes. Because approximately half of the meiotic cells at metaphase I are of each type, the ratio of the four possible types of gametes is 1:1:1:1.

ples in their own experiments; they later discovered Mendel’s paper and found that it explained their own research observations. By this time biologists had a much greater appreciation of the value of quantitative experimental methods. Correns gave credit to Mendel by naming the basic laws of inheritance after him. Although gametes and fertilization were known at the time Mendel carried out his research, mitosis and meiosis had not yet been discovered. It is truly remarkable that Mendel formulated his ideas mainly on the basis of mathematical abstractions. Today his principles are much easier to understand because we relate the transmission of genes to the behavior of chromosomes. The details of mitosis and meiosis were described during the late 19th century, and in 1902, American biologist Walter Sutton and German biologist Theodor Boveri independently pointed out the connection between Mendel’s segregation of alleles and the separation of homologous chromosomes during meiosis. This connection developed into the chromosome theory of inheritance, or the Sutton–Boveri theory, which stated that inheritance can be explained by assuming that genes are linearly arranged in specific locations along the chromosomes. The chromosome theory of inheritance was initially controversial, because at that time there was no direct evidence that genes are found on chromosomes. However, new research provided the findings necessary for wider acceptance and extension of these ideas and their implications. For example, the work of American geneticist Thomas Hunt Morgan in 1910 provided evidence for the location of a particular gene (white eye color) on a specific chromosome (the X chromosome) in fruit flies. Morgan and his graduate students also provided insight into the way genes are organized on chromosomes; we discuss some of Morgan’s research contributions later in the chapter.

Review ■ ■ ■

What are the relationships among loci, genes, and alleles? What is Mendel’s principle of segregation? What is Mendel’s principle of independent assortment?

11.2 USING PROBABILITY TO PREDICT MENDELIAN INHERITANCE ■ ■ LEARNING OBJECTIVE 5

Apply the product rule and sum rule appropriately when predicting the outcomes of genetic crosses.

All genetic ratios are properly expressed in terms of probabilities. In monohybrid crosses, the expected ratio of the dominant and recessive phenotypes is 3:1. The probability of an event is its expected frequency. Therefore, we can say there are 3 chances in 4 (or –34) that any particular individual offspring of two heterozygous individuals will express the dominant phenotype and 1 chance in 4 (or –14) that it will express the recessive phenotype. Although we sometimes speak in terms of percentages, probabilities are calcu-

Second toss

Probability is

First toss

Probability is

1 2

1

Probability is

1 2

1

2

2

1 2

Probability is

1 = 1

1

1 1 ✕ 2 = 4

1

✕

2

4

2

✕

2 ✕

1 2

1 = 1 2

4

1 = 1 2

4

FIGURE 11-10 The rules of probability For each coin toss, the probability of heads is –21 , and the probability of tails is also –21 . Because the outcome of the first toss is independent of the outcome of the second, the combined probabilities of the outcomes of successive tosses are calculated by multiplying their individual probabilities (accord1 ing to the product rule: –21 × –21 = – 4 ). These same rules of probability predict genetic events.

lated as fractions (such as –34) or decimal fractions (such as 0.75). If an event is certain to occur, its probability is 1; if it is certain not to occur, its probability is 0. A probability can be 0, 1, or some number between 0 and 1. The Punnett square lets you combine two or more probabilities. When you use a Punnett square, you are following two important statistical principles known as the product rule and the sum rule. The product rule predicts the combined probabilities of independent events. Events are independent if the occurrence of one does not affect the probability that the other will occur. For example, the probability of obtaining heads on the first toss of a coin is –12 ; the probability of obtaining heads on the second toss (an independent event) is also –12 . If two or more events are independent of each other, the probability of both occurring is the product of their individual probabilities. If this seems strange, keep in mind that when we multiply two numbers that are less than 1, the product is a smaller number. Therefore, the probability of obtaining heads two times in a row is –12 × –12 = –14 , or 1 chance in 4 (FIG. 11-10). Similarly, we can apply the product rule to genetic events. If both parents are Bb, what is the probability they will produce a child who is bb? For the child to be bb, he or she must receive a b gamete from each parent. The probability of a b egg is –12 , and the probability of a b sperm is also –12 . Like the outcomes of the coin tosses, these probabilities are independent, so we combine them by the product rule (–12 × –12 = –14). You might like to check this result using a Punnett square. The sum rule predicts the combined probabilities of mutually exclusive events. In some cases, there is more than one way to obtain a specific outcome. These different ways are mutually exclusive; if one occurs, the other(s) cannot. For example, if both parents are

In qu i ri n g A b o u t S O LV I N G G E N E T I C S P RO B L E M S How do you solve simple Mendelian genetics problems? Such problems can be fun and easy to work if you follow certain conventions and are methodical in your approach. 1. Always use standard designations for the generations. The generation in which a particular genetic experiment is begun is called the P, or parental, generation. Offspring of this generation are called the F1, or first filial, generation. The offspring resulting when two F1 individuals are bred constitute the F2, or second filial, generation. 2. Write down a key for the symbols you are using for the allelic variants of each locus. Use an uppercase letter to designate a dominant allele and a lowercase letter to designate a recessive allele. Use the same letter of the alphabet to designate both alleles of a particular locus. If you are not told which allele is dominant and which is recessive, the phenotype of the F1 generation is a good clue. 3. Determine the genotypes of the parents of each cross by using the following types of evidence: • Are they from true-breeding lines? If so, they should be homozygous. • Can their genotypes be reliably deduced from their phenotypes? This is usually true if they express the recessive phenotype. • Do the phenotypes of their offspring provide any information? Exactly how this is done is discussed shortly. 4. Indicate the possible kinds of gametes formed by each of the parents. It is helpful to draw a circle around the symbols for each kind of gamete. • If it is a monohybrid cross, we apply the principle of segregation; that is, a heterozygote Aa forms two kinds of gametes, A and a. A homozygote, such as aa, forms only one kind of gamete, a.

•

If it is a dihybrid cross, we apply the principles of segregation and independent assortment. For example, an individual heterozygous for two loci would have the genotype AaBb. Allele A segregates from a, and B segregates from b. The assortment of A and a into gametes is independent of the assortment of B and b. A is equally likely to end up in a gamete with B or b. The same is true for a. Thus, an individual with the genotype AaBb produces four kinds of gametes in equal amounts: AB, Ab, aB, and ab.

5. Set up a Punnett square, placing the possible types of gametes from one parent down the left side and the possible types from the other parent across the top. 6. Fill in the Punnett square. Avoid confusion by consistently placing the dominant allele first and the recessive allele second in heterozygotes (Aa, never aA). If it is a dihybrid cross, always write the two alleles of one locus first and the two alleles of the other locus second. It does not matter which locus you choose to write first, but once you have decided on the order, it is crucial to maintain it consistently. This means that if the individual is heterozygous for both loci, you will always use the form AaBb. Writing this particular genotype as aBbA, or even as BbAa, would cause confusion. 7. If you do not need to know the frequencies of all the expected genotypes and phenotypes, you can use the rules of probability as a shortcut instead of making a Punnett square. For example, if both parents are AaBb, what is the probability of an AABB offspring? To be AA, the offspring must receive an A gamete from each parent. The probability that a given gamete is A is –21 , and each gamete represents an independent event, so combine their probabilities by multiplying (–21 × –21 = –41 ). The probability

Bb, what is the probability that their first child will also have the Bb genotype? There are two different ways these parents can have a Bb child: either a B egg combines with a b sperm (probability –14 ), or a b egg combines with a B sperm (probability –14 ). Naturally, if there is more than one way to get a result, the chances of its being obtained improve; we therefore combine the probabilities of mutually exclusive events by summing (adding) their individual probabilities. The probability of obtaining a Bb child in our example is therefore –14 + –14 = –12 . (Because there is only

1 of BB is calculated similarly and is also – 4. The probability of AA is independent of the probability of BB, so again, use the product rule to obtain their combined 1 probabilities (–41 × –41 = — 16 ).

8. Very often the genotypes of the parents can be deduced from the phenotypes of their offspring. In peas, for example, the allele for yellow seeds (Y) is dominant to the allele for green seeds (y). Suppose plant breeders cross two plants with yellow seeds, but you don’t know whether the yellow-seeded plants are homozygous or heterozygous. You can designate the cross as: Y_ × Y_. The offspring of the cross are allowed to germinate and grow, and their seeds are examined. Of the offspring, 74 produce yellow seeds, and 26 produce green seeds. Knowing that green-seeded plants are recessive, yy, the answer is obvious: Each parent contributed a y allele to the greenseeded offspring, so the parents’ genotypes must have been Yy × Yy. Sometimes it is impossible to determine from the data given whether an individual is homozygous dominant or heterozygous. For example, suppose a cross between a yellow-seeded plant and a green-seeded plant yields offspring that all produce yellow seeds. From the information given about the parents, you can designate the cross as: Y_ × yy. Because all the offspring have yellow seeds, they have the genotype Yy. You can deduce this because each parent contributes one allele. (The offspring have yellow seeds, so they must have at least one Y allele; however, the green-seeded parent can contribute only a y allele to the offspring.) Therefore, the cross is most likely (YY × yy), but you cannot be absolutely certain about the genotype of the yellowseeded parent. Further crosses would be necessary to determine this.

one way these heterozygous parents can produce a homozygous recessive child, bb, that probability is only –14 . The probability of a homozygous dominant child, BB, is likewise –14 .)

The rules of probability can be applied to a variety of calculations The rules of probability have wide applications. For example, what are the probabilities that a family with two (and only two) children

will have two girls, two boys, or one girl and one boy? For purposes of discussion, let’s assume male and female births are equally probable. The probability of having a girl first is –12 , and the probability of having a girl second is also –12 . These are independent events, so we combine their probabilities by multiplying: –12 × –12 = –14 . Similarly, the probability of having two boys is –14 . In families with both a girl and a boy, the girl can be born first or the boy can be born first. The probability that a girl will be born first is –12 , and the probability that a boy will be born second is also 1 –2 . We use the product rule to combine the probabilities of these two independent events: –12 × –12 = –14 . Similarly, the probability that a boy will be born first and a girl second is –14 . These two kinds of families represent mutually exclusive outcomes, that is, two different ways of obtaining a family with one boy and one girl. Having two different ways of obtaining the desired result improves our chances, so we use the sum rule to combine the probabilities: –14 + –14 = –12 . In working with probabilities, keep in mind a point that many gamblers forget: Chance has no memory. If events are truly independent, past events have no influence on the probability of the occurrence of future events. Probability only has predictive value in the long run, over many trials. (Recall that Mendel counted hundreds of offspring for each cross, and this was one of the reasons for his success.) When working probability problems, common sense is more important than blindly memorizing rules. Examine your results to see whether they appear reasonable; if they don’t, re-evaluate your assumptions. (See Inquiring About: Solving Genetics Problems for procedures to solve genetics problems, including when to use the rules of probability.)

Review ■

■

Use the rules of probability to answer the following question: In a cross between homozygous pea plants with yellow, round seeds (YYRR) and homozygous pea plants with green, wrinkled seeds (yyrr), what is the probability of an F2 plant having yellow, round seeds? In the previous question, did you use the product rule or the sum rule? Why?

11.3 INHERITANCE AND CHROMOSOMES ■ ■ LEARNING OBJECTIVES 6 Define linkage and relate it to specific events in meiosis. 7 Show how data from a test cross involving alleles of two loci can be used 8

to distinguish between independent assortment and linkage. Discuss the genetic determination of sex and the inheritance of X-linked genes in mammals.

It is a measure of Mendel’s genius that he worked out the principles of segregation and independent assortment without knowing anything about meiosis or the chromosome theory of inheritance. The chromosome theory of inheritance also helps explain certain apparent exceptions to Mendelian inheritance. One of these socalled exceptions involves linked genes.

Linked genes do not assort independently Beginning around 1910, the research of geneticist Thomas Hunt Morgan and his graduate students extended the concept of the chromosome theory of inheritance. Morgan’s research organism was the fruit fly (Drosophila melanogaster). Just as the garden pea was an excellent model research organism for Mendel’s studies, the fruit fly was perfect for extending general knowledge about inheritance. Fruit flies have a short life cycle—just 14 days—and their small size means that thousands can be kept in a research lab. The large number of individuals increases the chance of identifying mutants. Also, fruit flies have just four pairs of chromosomes, one of which is a pair of sex chromosomes. By carefully analyzing the results of crosses involving fruit flies, Morgan and his students demonstrated that genes are arranged in a linear order on each chromosome. Morgan also showed that independent assortment does not apply if the two loci lie close together in the same pair of homologous chromosomes. In fruit flies there is a locus controlling wing shape: the dominant allele V for normal wings and the recessive allele v for abnormally short, or vestigial, wings. Another locus controls body color: the dominant allele B for gray body and the recessive allele b for black body. If a homozygous BBVV fly is crossed with a homozygous bbvv fly, the F1 flies all have gray bodies and normal wings, and their genotype is BbVv. Because these loci happen to lie close to each other in the same pair of homologous chromosomes, their alleles do not assort independently; instead, they are linked genes that tend to be inherited together. Linkage is the tendency for a group of genes on the same chromosome to be inherited together in successive generations. You can readily observe linkage in the results of a test cross in which heterozygous F1 flies (BbVv) are mated with homozygous recessive (bbvv) flies (FIG. 11-11). Because heterozygous individuals are mated to homozygous recessive individuals, this test cross is similar to the test cross described earlier. However, it is called a two-point test cross because alleles of two loci are involved. If the loci governing these traits were unlinked—that is, on different chromosomes—the heterozygous parent in a test cross would produce four kinds of gametes (BV, Bv, bV, and bv) in equal numbers. This independent assortment would produce offspring with new gene combinations not present in the parental generation. Recall that any process that leads to new allele combinations is called genetic recombination. In our example, gametes Bv and bV are recombinant types. The other two kinds of gametes, BV and bv, are parental types because they are identical to the gametes produced by the P generation. Of course, the homozygous recessive parent produces only one kind of gamete, bv. Thus, if independent assortment were to occur in the F1 flies, approximately 25% of the test-cross offspring would be gray-bodied and normal-winged (BbVv), 25% black-bodied and normal-winged (bbVv), 25% gray-bodied and vestigial-winged (Bbvv), and 25% black-bodied and vestigial-winged (bbvv). Notice that the two-point test cross lets us determine the genotypes of the offspring directly from their phenotypes. By contrast, the alleles of the loci in our example do not undergo independent assortment because they are linked. Alleles at different loci but close to one another on a given chromosome

KEY EXPERIMENT QUESTION: How can linkage be recognized in fruit flies? HYPOTHESIS: Linkage can be recognized when an excess of parental-type offspring and a deficiency of recombinant-type offspring are produced in a two-point test cross. EXPERIMENT: Fruit flies with gray, normal wings (BbVv) are crossed with flies that have black, vestigial wings (bbvv). If the alleles for color and wing shape are not linked (i.e., the alleles assort independently), the offspring will consist of an equal number of each of four phenotypes (bottom row).

Calculating the frequency of crossingover reveals the linear order of linked genes on a chromosome

Gray, normal wings BbVv

Parental-type gametes

Recombinant-type gametes

BV

bv

Bv

bV

Gray, normal

Black, vestigial

Gray, vestigial

Black, normal

BbVv

bbvv

Bbvv

bbVv

Expected results, independent assortment

575

575

575

575

Actual results

965

944

206

185

bv

Black, vestigial wings bbvv

RESULTS AND CONCLUSION: In the 2300 offspring (bottom row) of an actual cross, about 1909 of the offspring belong to each of the two parental types (83% total), and 391 belong to each of the two recombinant types (17% total). Thus, loci for wing length and body color are linked on a homologous chromosome pair.

FIGURE 11-11 A two-point test cross to detect linkage in fruit flies

tend to be inherited together; because chromosomes pair and separate during meiosis as units, the alleles at different loci on a given chromosome tend to be inherited as units. If linkage were complete, only parental-type flies would be produced, with approximately 50% having gray bodies and normal wings (BbVv) and 50% having black bodies and vestigial wings (bbvv). However, in our example, the offspring also include some gray-bodied, vestigial-winged flies and some black-bodied, normal-winged flies. These are recombinant flies, having received a recombinant-type gamete from the heterozygous F1 parent. Each recombinant-type gamete

TABLE 11-3

arose by crossing-over between these loci in a meiotic cell of a heterozygous female fly. (Fruit flies are unusual in that crossing-over occurs only in females and not in males; it is far more common for crossing-over to occur in both sexes of a species.) When chromosomes pair and undergo synapsis, crossing-over occurs as homologous (nonsister) chromatids exchange segments of chromosome material by a process of breakage and rejoining catalyzed by enzymes (FIG. 11-12). Also see the discussion of prophase I in Chapter 10, Section 10.4.

In our fruit fly example (see Fig. 11-11), 391 of the offspring are recombinant types: gray flies with vestigial wings, Bbvv (206 of the total); and black flies with normal wings, bbVv (185 of the total). The remaining 1909 offspring are parental types. These data can be used to calculate the percentage of crossing-over between the loci (TABLE 11-3). You do this by adding the number of individuals in the two recombinant types of offspring, dividing by the total number of offspring, and multiplying by 100: 391 ÷ 2300 = 0.17; 0.17 × 100 = 17%. Thus, the V locus and the B locus have 17% recombination between them. During a single meiotic division, crossing-over may occur at several different points along the length of each homologous chromosome pair. In general, crossing-over is more likely to occur between two loci if they lie far apart on the chromosome and less likely to occur if they lie close together. Because of this rough correlation between the frequency of recombination of two loci and the linear distance between them, a genetic map of the chromosome can be generated by converting the percentage of recombination to map units. By convention, 1% recombination between two loci equals a distance of 1 map unit, so the loci in our example are 17 map units apart.

How to Determine Recombination Frequency from a Two-Point Test Cross Test-Cross Results (from Fig. 11-11)

Type of Offspring

Parental Type

Recombinant Type

Phenotype

Gray, normal

Black, vestigial

Gray, vestigial

Black, normal

Genotype

BbVv

bbvv

Bbvv

bbVv

Number of offspring

965

944

206

185

Calculations of recombination frequency

1. Number of parental-type offspring = 1909 2. Number of recombinant-type offspring = 391 3. Total number of offspring = 1909 + 391 = 2300 4. Recombination frequency = 391/2300 × 100 = 17%

KEY POINT

The exchange of segments between chromatids of homologous chromosomes is the mechanism of recombination of linked genes.

v

v

B

B

b

b

V

v

B

b

Crossing-over between a pair of homologous (nonsister) chromatids

5 map units

3 map units

A

2 map units 3 map units

B

C

(a) If the recombination between A and C is 8% (8 map units), then B must be in the middle.

0 1 2 3 4 5 6 7 8

V

0 1 2 3 4 5 6 7 8

V

Two homologous chromosomes undergo synapsis in meiosis

A C

B

(b) If the recombination between A and C is 2%, then C must be in the middle.

Meiosis I

FIGURE 11-13 Gene mapping

V

V

B

b

Meiosis II

v

v

B

b

Gene order (that is, which locus lies between the other two) is determined by the percentage of recombination between each of the possible pairs. In this hypothetical example, the percentage of recombination between locus A and locus B is 5% (corresponding to 5 map units) and that between B and C is 3% (3 map units). There are two alternatives for the linear order of these alleles.

V

V

v

v

B

b

B

b

Parental type

Recombinant type

Recombinant type

many plants, especially those that are important crops. In addition, researchers have used genetic methods to develop a detailed map for Escherichia coli, a bacterium with a single, circular DNA molecule, and many other prokaryotes and viruses. They have made much more sophisticated maps of chromosomes of various species by means of recombinant DNA technology (discussed in Chapter 15). Using these techniques, the Human Genome Project has produced detailed maps of human chromosomes (discussed in Chapter 16).

Parental type

Four haploid cells produced

FIGURE 11-12 Animated Crossing-over Genes located far apart on a chromosome have a greater probability of being separated by crossing-over than do genes that are closer together.

Scientists have determined the frequencies of recombination between specific linked loci in many species. All the experimental results are consistent with the hypothesis that genes are present in a linear order in the chromosomes. FIGURE 11-13 illustrates the traditional method for determining the linear order of genes in a chromosome. By putting together the results of numerous crosses, scientists have developed detailed linkage maps for many eukaryotes, including the fruit fly, the mouse, yeast, Neurospora (a fungus), and

Sex is generally determined by sex chromosomes In some species, environmental factors exert a large control over an individual’s sex. However, genes are the most important sex determinants in most eukaryotic organisms. The major sex-determining genes of mammals, birds, and many insects are carried by sex chromosomes. Typically, members of one sex have a pair of similar sex chromosomes and produce gametes that are all identical in sex chromosome constitution. The members of the other sex have two different sex chromosomes and produce two kinds of gametes, each bearing a single kind of sex chromosome. The cells of female mammals, including humans, contain two X chromosomes. In contrast, the males have a single X chromosome and a smaller Y chromosome that bears only a few active genes (FIG. 11-14). For example, human females have 22 pairs of autosomes, which are chromosomes other than the sex chromosomes, plus a pair of X chromosomes; males have 22 pairs of autosomes plus one X chromosome and one Y chromosome. Domestic

© Biophoto Associates/ Photo Researchers, Inc.

2 μm

FIGURE 11-14 SEM of human Y chromosome (left) and X chromosome (right) Each chromosome is in the duplicated state and consists of two identical chromatids.

cats have 19 pairs of autosomes, to which are added a pair of X chromosomes in females, or an X plus a Y in males. In contrast, sex determination is based on different chromosome systems in animals other than mammals (TABLE 11-4). The Y chromosome determines male sex in most species of mammals Do male humans have a male phenotype because they have only one X chromosome, or because they have a Y chromosome? Much of the traditional evidence bearing on this question comes from studies of people with abnormal sex chromosome constitutions (see discussion in Chapter 16). A person with an XXY constitution is a nearly normal male in external appearance, although his testes are underdeveloped (Klinefelter syndrome). A person with one X but no Y chromosome has the overall appearance of a female but has defects such as short stature and undeveloped ovaries (Turner syndrome). An embryo with a Y but no X does not survive. Based on these and other obser-

TABLE 11-4

vations, biologists concluded that all individuals require at least one X, and the Y is the male-determining chromosome. Geneticists have identified several genes on the Y chromosome that are involved in determination of the male sex. The sex reversal on Y (SRY) gene, the major male-determining gene on the Y chromosome, acts as a “genetic switch” that causes testes to develop in the fetus. The developing testes then secrete the hormone testosterone, which causes other male characteristics to develop. Several other genes on the Y chromosome also play a role in sex determination, as do many genes on the X chromosome, which explains why an XXY individual does not have a completely normal male phenotype. Some genes on the autosomes also affect sex development. Evidence suggests that the X and Y chromosomes of mammals originated as a homologous pair of autosomes. During the evolution of the X and Y chromosomes, almost all of the original functional genes were retained on the X chromosome and lost from the Y chromosome. Today, about 95% of the Y chromosome is male-specific. Either the Y chromosome retained certain genes that coded for proteins that determine maleness, or perhaps mutations occurred in existing genes on the Y chromosome that made it the male-determining chromosome. Thus, the sex chromosomes are not truly homologous in their present forms because they are not similar in size, shape, or genetic constitution. Nevertheless, the Y chromosome has short, homologous “pairing regions” at its tips that lets it synapse and exchange genetic material with the X chromosome during meiosis. Half the sperm contain an X chromosome, and half contain a Y chromosome. All normal eggs bear a single X chromosome. Fertilization of an X-bearing egg by an X-bearing sperm results in an XX (female) zygote, or fertilized egg; fertilization by a Y-bearing sperm results in an XY (male) zygote. You might expect to have equal numbers of X- and Y-bearing sperm and a 1:1 ratio of females to males. However, in humans

Representative Examples of Sex Determination in Animals

X-Y system Mammals, including humans

X-O system Certain insects

Z-W system Birds, fishes, and some insects

Haplo-diploid system Most bees and ants

Male XY Occurs when sperm cell contains a Y chromosome

Male XO Occurs when sperm cell contains no sex chromosome

Male ZW Occurs when an egg cell contains a W chromosome

Male Haploid No sex chromosomes; male develops from unfertilized egg

Female XX Occurs when sperm cell contains an X chromosome

Female XX Occurs when sperm cell contains an X chromosome

Female ZZ Occurs when an egg cell contains a Z chromosome

Female Diploid No sex chromosomes; female develops from fertilized egg

more males are conceived than females, and more males die before birth. Even at birth the ratio is not 1:1; about 106 boys are born for every 100 girls. The Y-bearing sperm appear to have a selective advantage, possibly because sperm containing the Y chromosome are smaller and have less mass than sperm containing the X chromosome; it is hypothesized that, on average, the sperm carrying the Y chromosome can swim slightly faster than X-bearing sperm to reach the egg. X-linked genes have unusual inheritance patterns The human X chromosome contains many loci that are required in both sexes. Genes located in the X chromosome, such as those governing color perception and blood clotting, are sometimes called sex-linked genes. It is more appropriate, however, to refer to them as X-linked genes because they follow the transmission pattern of the X chromosome and, strictly speaking, are not linked to the sex of the organism. A female receives one X from her mother and one X from her father. A male receives his Y chromosome, which makes him male, from his father. From his mother he inherits a single X chromosome and therefore all his X-linked genes. In the male, every allele present on the X chromosome is expressed, whether that allele was dominant or recessive in the female parent. A male is neither homozygous nor heterozygous for his Xlinked alleles; instead, he is always hemizygous—that is, he has Color-blind male Xe Y

Heterozygous female XE Xe

XE

Gametes

Normal male XE Y

Xe

Gametes Xe

only one copy of each X-linked gene. The significance of hemizygosity in males, including human males, is that rare, recessive Xlinked genes are expressed, making males more likely to be affected by the many genetic disorders associated with the X chromosome. In comparison only a few genes are located exclusively on the Y chromosome. X and Y chromosomes also have short areas where they carry the same genes, enabling the X and Y to pair during meiosis I. We will use a simple system of notation for problems involving X linkage, indicating the X chromosome and incorporating specific alleles as superscripts. For example, the symbol Xe signifies a recessive X-linked allele for color blindness and XE the dominant X-linked allele for normal color vision. The Y chromosome is written without superscripts because it does not carry the locus of interest. Two recessive X-linked alleles must be present in a female for the abnormal phenotype to be expressed (XeXe), whereas in the hemizygous male a single recessive allele (Xe Y) causes the abnormal phenotype. As a consequence, these abnormal alleles are much more frequently expressed in male offspring. A heterozygous female may be a carrier, an individual who possesses one copy of a mutant recessive allele but does not express it in the phenotype (XEXe). To be expressed in a female, a recessive X-linked allele must be inherited from both parents. A color-blind female, for example, must have a color-blind father and a mother who is homozygous or heterozygous for a recessive allele for color blindness (FIG. 11-15). Heterozygous female XE Xe

XE

Gametes

Gametes Normal female XE Xe

Color-blind female Xe Xe

XE

Normal female XE XE

Carrier Y

Xe

Normal male XE Y

Normal female XE Xe Carrier

Color-blind male Xe Y

Y

Normal male XE Y

Color-blind male Xe Y

Normal color vision Color-blind

(a) To be color-blind, a female must inherit alleles for color blindness from both parents.

FIGURE 11-15 Animated X-linked red–green color blindness Note that the Y chromosome does not carry a gene for color vision.

(b) If a normal male mates with a carrier (heterozygous) female, half of their sons would be expected to be color-blind and half of their daughters would be expected to be carriers.

The homozygous combination is unusual because the frequency of alleles for color blindness is relatively low. In contrast, a colorblind male need only have a mother who is heterozygous for color blindness; his father can have normal vision. Therefore, X-linked recessive traits are generally much more common in males than in females, a fact that may partially explain why human male embryos are more likely to die.

KEY POINT

Dosage compensation equalizes the expression of X-linked genes in males and females The X chromosome contains numerous genes required by both sexes, yet a normal female has two copies (“doses”) for each locus, whereas a normal male has only one. Dosage compensation is a mechanism that makes equivalent the two doses in the female and the single dose in the male. Because of dosage compensation,

In female mammals, one of the two X chromosomes is randomly inactivated during early development, equalizing the level of expression of genes at loci on the X chromosome in males and females. 1

1

The zygote and early embryonic cells have two X chromosomes, one from each parent.

2

Random inactivation of one X chromosome occurs early in development. Roughly half the cells inactivate one X chromosome (left cell ), and the other half inactivate the second X chromosome (right cell ). The inactive X chromosome is visible as a Barr body near the nuclear envelope.

3

Chromosome inactivation persists through subsequent mitotic divisions, resulting in patches of cells in the adult body.

Nucleus

2

2

Barr body (inactivated chromosome)

3

3

Eunice Percy/Animals Animals

(a) Inactivation of one X chromosome in female cells.

FIGURE 11-16 Animated Dosage compensation in female mammals

(b) Persian calico cat. A calico cat has X-linked genes for both black and yellow (or orange) fur. Because of random X chromosome inactivation, black is expressed in some groups of cells and orange is expressed in others. (The patches of white fur are due to the presence of other genes that affect fur color.)

males and females produce the same amounts of proteins coded by X-linked genes. Male fruit flies accomplish this by making their single X chromosome more active. In most fly tissues, the metabolic activity of a single male X chromosome is equal to the combined metabolic activity of the two X chromosomes present in the female. Our understanding of dosage compensation in humans and other mammals is incomplete, but the process generally involves the random inactivation of one of the two X chromosomes in the female (FIG. 11-16a). During interphase, a dark spot of chromatin is visible at the edge of the nucleus of each female mammalian cell when stained and observed with a microscope. This dark spot, a Barr body, is a dense, metabolically inactive X chromosome. The other X chromosome resembles the metabolically active autosomes; during interphase, it is a greatly extended thread that is not evident in light microscopy. From this and other evidence, the British geneticist Mary Lyon hypothesized in 1961 that in most cells of a female mammal, only one of the two X chromosomes is active; the other is inactive and is condensed as a Barr body. Actually, however, X chromosome inactivation is never complete; as many as 25% of the genes on the inactive chromosome are expressed to some degree. X chromosome inactivation is a random event in each somatic (body) cell of the female embryo. A female mammal that is heterozygous at an X-linked locus expresses one of the alleles in about half her cells, and the other allele in the other half. X chromosome inactivation is sometimes evident in the phenotype. Mice and cats have several X-linked genes for certain coat colors. Females that are heterozygous for such genes may show patches of one coat color in the middle of areas of the other coat color. This phenomenon, termed variegation, is evident in tortoiseshell and calico cats (FIG. 11-16b). Early in development, when relatively few cells are present, X chromosome inactivation occurs randomly in each cell. X inactivation is then maintained during all subsequent divisions of that cell line. When any one of these cells divides by mitosis, the cells of the resulting clone (a group of genetically identical cells) all have the same active X chromosome, and therefore a patch develops of cells that all express the same color. You might wonder why variegation is not always apparent in females heterozygous at X-linked loci. The answer is that, although variegation usually occurs, we may need to use special techniques to observe it. For example, color blindness is caused by a defect involving the pigments in the cone cells in the retina of the eye. In at least one type of red–green color blindness, the retina of a heterozygous female actually contains patches of abnormal cones, but the patches of normal cones are enough to provide normal color vision. Variegation can be very hard to observe in cases where cell products become mixed in bodily fluids. For instance, in females heterozygous for the allele that causes hemophilia, only half the cells responsible for producing a specific blood-clotting factor do so, but they produce enough to ensure that the blood clots normally.

Review ■ ■

What are linked genes? How do geneticists use a two-point test cross to detect linkage?

■

■

Which chromosome determines the male sex in humans and other mammals? What is dosage compensation, and what does it generally involve in mammals?

11.4 EXTENSIONS OF MENDELIAN GENETICS ■ ■ LEARNING OBJECTIVES 9 Explain some of the ways genes may interact to affect the phenotype; discuss how a single gene can affect many features of the organism simultaneously. 10 Distinguish among incomplete dominance, codominance, multiple alleles, epistasis, and polygenic inheritance. 11 Describe norm of reaction and give an example.

The relationship between a given locus and the trait it controls may or may not be simple. A single pair of alleles of a locus may regulate the appearance of a single trait (such as tall versus short in garden peas). Alternatively, a pair of alleles may participate in the control of several traits, or alleles of many loci may interact to affect the phenotypic expression of a single character. Not surprisingly, these more complex relationships are quite common. You can assess the phenotype on one or many levels. It may be a morphological trait, such as shape, size, or color. It may be a physiological trait or even a biochemical trait, such as the presence or absence of a specific enzyme required for the metabolism of some specific molecule. In addition, changes in the environmental conditions under which the organism develops may alter the phenotypic expression of genes.

Dominance is not always complete Studies of the inheritance of many traits in a wide variety of organisms have shown that one member of a pair of alleles may not be completely dominant over the other. In such instances, it is inaccurate to use the terms dominant and recessive. The plants commonly called four o’clocks (Mirabilis jalapa) may have red or white flowers. Each color breeds true when these plants are self-pollinated. What flower color might you expect in the offspring of a cross between a red-flowering plant and a whiteflowering one? Without knowing which is dominant, you might predict that all would have red flowers or all would have white flowers. German botanist Carl Correns, one of the rediscoverers of Mendel’s work, first performed this cross and found that all F1 offspring have pink flowers! Does this result in any way indicate that the principles of inheritance that Mendel deduced are wrong? Did the parental traits blend inseparably in the offspring? Quite the contrary, for when two of these pink-flowered plants are crossed, red-flowered, pinkflowered, and white-flowered offspring appear in a ratio of 1:2:1 (FIG. 11-17). In this instance, as in all other aspects of the scientific process, results that differ from those hypothesized prompt scientists to re-examine and modify their hypotheses to account for the exceptional results. The pink-flowered plants are clearly the heterozygous individuals, and neither the red allele nor the white

KEY POINT

TABLE 11-5

In incomplete dominance, an F1 heterozygote has a phenotype intermediate between its parents. Red

White

P generation R1R1

R 2R 2 Pink

F1 generation

Gametes Gametes

A B AB O

Genotype

Surface Antigen on RBC

Serum Antibodies to A or B Antigens

IAIA or IAi IBIB or IBi I AI B ii

A B A, B None

Anti-B Anti-A None Anti-A, anti-B

*This table and the discussion of the ABO system have been simplified somewhat. Note that the body produces antibodies against the antigens lacking on its own red blood cells (RBCs). Because of their specificity for the corresponding antigens, these antibodies are used in standard tests to determine blood types.

All R 1R 2

R1

Phenotype (blood type)

ABO Blood Types

R2

R1R1

R1R2

R1R 2

R 2R 2

R1

R2

F2 generation

FIGURE 11-17 Animated Incomplete dominance in four

o’clocks Two incompletely dominant alleles, R1 and R2, are responsible for red, white, and pink flower colors. Red-flowered plants are R1R1, whiteflowered plants are R2R2, and heterozygotes (R1R2) are pink. Note that uppercase letters are used for both alleles because neither is recessive to the other.

allele is completely dominant. When the heterozygote has a phenotype intermediate between those of its two parents, the genes show incomplete dominance. In these crosses, the genotypic and phenotypic ratios are identical. Incomplete dominance is not unique to four o’clocks, and additional examples of incomplete dominance are known in both plants and animals. For example, true-breeding white chickens and true-breeding black chickens produce bluish gray offspring, or Andalusian blues, when crossed. In both cattle and horses, reddish coat color is not completely dominant to white coat color. Heterozygous individuals have a mixture of reddish hairs and white hairs, which is called roan. If you saw a white mare nursing a roan foal, what would you guess was the coat color of the foal’s father? Because the reddish and white colors are expressed independently (hair by hair) in the roan heterozygote, scientists sometimes refer to this as a case of codominance. Strictly speaking, the term incomplete dominance refers to instances in which the heterozygote is intermediate in phenotype, and codominance refers to instances in which the heterozygote simultaneously expresses the phenotypes of both types of homozygotes.

Humans have four blood types (A, B, AB, and O), collectively called the ABO blood group. The human ABO blood group is an excellent example of codominant alleles. Blood types A, B, AB, and O are controlled by three alleles representing a single locus (TABLE 11-5). Allele IA codes for the synthesis of a specific glycoprotein, antigen A, which is expressed on the surface of red blood cells. (Immune responses are discussed in Chapter 45.) Antigens are substances capable of stimulating an immune response. Allele IB leads to the production of antigen B, a different but related glycoprotein. Allele i is allelic to IA and IB, but it does not code for an antigen. Individuals with the genotype IAIA or IAi have blood type A. People with genotype IBIB or IBi have blood type B. Those with genotype IAIB have blood type AB, whereas those with genotype ii have blood type O. These results show that neither allele IA nor allele IB is dominant to the other. Both alleles are expressed phenotypically in the heterozygote and are therefore codominant to each other, although each is dominant to allele i. At one time, determining blood types was used to settle cases of disputed parentage. Although blood-type tests can exclude someone as a possible parent of a particular child, they can never prove a certain person is the parent; they only determine he or she could be. Could a man with blood type AB be the father of a child with blood type O? Could a woman with blood type O be the mother of a child with blood type AB? Could a type B child with a type A mother have a type A father or a type O father?3

Multiple alleles for a locus may exist in a population Most of our examples so far have dealt with situations in which each locus was represented by a maximum of two allelic variants (three in the example of blood types). It is true that a single diploid individual has a maximum of two different alleles for a particular locus and that a haploid gamete has only one allele for each locus. However, if you survey a population, you may find more than two alleles for a particular locus, as you saw with the ABO blood group (see Table 11-5). If three or more alleles for a given locus exist within the population, that locus has multiple alleles. Research has shown that many loci have multiple alleles. Some alleles can be identified by the activity of a certain enzyme or by

3

The answer to all these questions is no.

some other biochemical feature but do not produce an obvious phenotype. Others produce a readily recognizable phenotype, and certain patterns of dominance can be discerned when the alleles are combined in various ways. In other series of multiple alleles, certain alleles may be codominant and others incompletely dominant. In such cases, the heterozygotes commonly have phenotypes intermediate between those of their parents.

A single gene may affect multiple aspects of the phenotype In our examples, the relationship between a gene and its phenotype has been direct, precise, and exact, and the loci have controlled the appearance of single traits. However, the relationship of gene to trait may not have such a simple genetic basis. Most genes affect several different characters. The ability of a single gene to have multiple effects is known as pleiotropy. Most cases of pleiotropy can be traced to a single fundamental cause. For example, a defective enzyme may affect the functioning of many types of cells. Pleiotropy is evident in many genetic diseases in which a single pair of alleles causes multiple symptoms. For example, people who are homozygous for the recessive allele that causes cystic fibrosis produce abnormally thick mucus in many parts of the body, including the respiratory, digestive, and reproductive systems (cystic fibrosis is discussed in Chapter 16).

Alleles of different loci may interact to produce a phenotype Several pairs of alleles may interact to affect a single phenotype, or one pair may inhibit or reverse the effect of another pair. One example of gene interaction is illustrated by the inheritance of combs in chickens, where two genes may interact to produce a novel phenotype (FIG. 11-18). The allele for a rose comb, R, is dominant to that for a single comb, r. A second, unlinked pair of alleles governs the inheritance of a pea comb, P, versus a single comb, p. A single-comb chicken is homozygous for the recessive allele at both loci (pprr). A rose comb chicken is either ppRR or ppRr, and a pea comb chicken is either PPrr or Pprr. When an R and P occur in the same individual, the phenotype is neither pea nor rose comb but a completely different type, a walnut comb. The walnut comb phenotype is produced whenever a chicken has one or two R alleles, plus one or two P alleles (that is, PPRR, PpRR, PPRr, or PpRr). What would you hypothesize about the types of combs among the offspring of two heterozygous walnut comb chickens, PpRr? How does this form of gene interaction affect the ratio of phenotypes in the F2 generation? Is it the typical Mendelian 9:3:3:1 ratio?4 Epistasis is a common type of gene interaction in which the presence of certain alleles of one locus can prevent or mask the expression of alleles of a different locus and express their own pheno-

4

The offspring of two heterozygous walnut comb chickens will have four genotypes in what appears to be a Mendelian 9:3:3:1 ratio: 9 walnut comb, 3 pea comb, 3 rose comb, and 1 single comb. This example is not a typical Mendelian 9:3:3:1 ratio because it involves a single character (comb shape) coded by alleles at two different loci. In Mendelian inheritance, the 9:3:3:1 ratio involves two characters (such as seed color and seed shape) coded by alleles at two loci.

Walnut comb PPRR, PpRR, PPRr, or PpRr

Pea comb PPrr or Pprr

Rose comb ppRR or ppRr

Single comb pprr

FIGURE 11-18 Animated Gene interaction in chickens Two gene pairs govern four chicken comb phenotypes. Chickens with walnut combs have the genotype P_R_. Chickens with pea combs have the genotype P_rr, and those with rose combs have the genotype ppR_. Chickens that are homozygous recessive for both loci, pprr, have a single comb. (The blanks represent either dominant or recessive alleles.)

type instead. (The term epistasis means “standing on.”) Unlike the chicken example, no novel phenotypes are produced in epistasis. Coat color in Labrador retrievers is an example of epistasis that involves a gene for pigment and a gene for depositing color in the coat (FIG. 11-19). The two alleles for the pigment gene are B for black coat and its recessive counterpart, b, for brown coat. The gene for depositing color in the coat has two alleles, E for the expression of black and brown coats and e, which is epistatic and blocks the expression of the B/b gene. The epistatic allele is recessive and therefore is expressed as a yellow coat only in the homozygous condition (ee), regardless of the combination of B and b alleles in the genotype.

In polygenic inheritance, the offspring exhibit a continuous variation in phenotypes Many human characters—such as height, body form, and skin pigmentation—are not inherited through alleles at a single locus. The same holds true for many commercially important characters in domestic plants and animals, such as milk and egg production. Alleles at several, perhaps many, loci affect each character. The term polygenic inheritance is applied when multiple independent pairs of genes have similar and additive effects on the same character. As many as 60 loci account for the inheritance of skin pigmentation in humans. To keep things simple in this example, we

Renee Stockdale/Animals Animals

Black BBEE, BbEE, BBEe or BbEe

Yellow BBee, Bbee or bbee

Chocolate bbEE or bbEe

FIGURE 11-19 Animated Epistasis in Labrador retrievers

depending on the level of aluminum in the soil before the flowers begin to develop. It is easy to manipulate the level of aluminum by adding alum or aluminum sulfate to the soil or by lowering the soil pH (in acidic soils, aluminum is more soluble). Under these conditions, hydrangea flowers are blue. In alkaline soils (made by adding limestone to the soil), aluminum is less soluble, and the flowers are pink (FIG. 11-21). Another example of the effect of environment on gene expression involves Himalayan rabbits. The phenotype of these rabbits is white fur except for dark patches on the ears, nose, and paws. The local surface temperature of a rabbit’s ears, nose, and paws is colder in the rabbit’s natural environment, and this temperature difference causes the production of dark fur. If you raise rabbits with the Himalayan genotype at a warm temperature

Two gene pairs interact to govern coat color in Labrador retrievers. Black Labs have the genotype B_E_; yellow Labs have the genotype B_ee or bbee; and chocolate Labs have the genotype bbE_. (The blanks represent either dominant or recessive alleles.)

X AaBbCc

Genes interact with the environment to shape phenotype Hydrangeas are shrubs grown for their attractive flowers. The color of certain hydrangea flowers ranges from blue to purple to pink

AaBbCc

20

15 Number of individuals (F2)

illustrate the principle of polygenic inheritance in human skin pigmentation with pairs of alleles at three unlinked loci. These can be designated A and a, B and b, and C and c. The capital letters represent incompletely dominant alleles producing dark skin. The more capital letters, the darker the skin because the alleles affect skin pigmentation in an additive fashion. A person with the darkest skin possible would have the genotype AABBCC, and a person with the lightest skin possible would have the genotype aabbcc. The F1 offspring of an aabbcc person and an AABBCC person are all AaBbCc and have an intermediate skin color. The F2 offspring of two such triple heterozygotes (AaBbCc × AaBbCc) would have skin pigmentation ranging from very dark to very light (FIG. 11-20). Polygenic inheritance is characterized by an F1 generation that is intermediate between the two completely homozygous parents and by an F2 generation that shows wide variation between the two parental types. When the number of individuals in a population is plotted against the amount of skin pigmentation and the points are connected, the result is a bell-shaped curve, a normal distribution curve. Most of the F2 generation individuals have one of the intermediate phenotypes; only a few show the extreme phenotypes of the original P generation (that is, the grandparents). On average, only 1 of 64 is as dark as the very dark grandparent, and only 1 of 64 is as light as the very light grandparent. The alleles A, B, and C each produce about the same amount of darkening of the skin; hence, the genotypes AaBbCc , AABbcc, AAbbCc, AaBBcc, aaBBCc, AabbCC, and aaBbCC all produce similar intermediate phenotypes.

6

1

Very dark

Skin pigmentation

Very light

FIGURE 11-20 Polygenic inheritance in human skin pigmentation This simplified example assumes that skin pigmentation in humans is governed by alleles of three unlinked loci. The alleles producing dark skin (A, B, and C) are represented by capital letters, but they are not dominant. Instead, their effects are additive. The number of dark dots, each signifying an allele producing dark skin, is counted to determine the phenotype. A wide range of phenotypes is possible when individuals of intermediate phenotype mate and have offspring (AaBbCc × AaBbCc). The expected distribution of phenotypes is consistent with the superimposed normal distribution curve.

© Wildscape/Alamy

FIGURE 11-21 Influence of the environment on hydrangea flower color Hydrangea flowers vary in color depending on availability of aluminum, which in turn is dependent on soil pH. Blue flowers develop in a soil pH of 5.5 or less, purple flowers in a soil pH of 5.6 to 6.4, and pink flowers in a soil pH of 6.5 to 7.0.

(30°C), the rabbits are completely white, with no dark patches on the ears, nose, or paws (FIG. 11-22a). If you raise Himalayan rabbits at a cooler temperature (25°C), however, they develop the characteristic dark patches of fur (FIG. 11-22b). Thus, genes can function differently in different environments. Now let’s examine a human example—height—in the context of genes and the environment. The inheritance of height in humans is polygenic and involves alleles representing ten or more loci. Because many genes are involved and because height is modified by a variety of environmental conditions, such as diet and general health, the height of most adults ranges from 4 ft 2 in (1.25 m) to 7 ft 2 in (2.15 m). The genes that affect height set limits for the phenotype—no human is 12 ft tall, for example—but the environment shapes the phenotype within its genetic limits. The range of phenotypic possibilities that can develop from a single genotype under different environmental conditions is known as the norm of reaction. In certain genotypes the norm of reaction is quite limited. In other genotypes, such as those involved in human height, the norm of reaction is quite broad. Although interactions between genes and the environment influence the phenotypes of many characters, it is difficult to de-

termine the exact contributions of genes and the environment to a given phenotype. In some cases the environment regulates the activity of certain genes, turning them on in some environmental conditions and off in others. Investigators argued for years about

(a) The Himalayan genotype at warm temperatures.

(b) The Himalayan genotype at cold temperatures.

FIGURE 11-22 Animated Influence of the environment on rabbit coat color These rabbits have the same genotype for coat color, but have been exposed to different temperatures.

the relative importance of nature (genetics/biology) versus nurture (environmental influences) regarding such human characters as intelligence, depression, bipolar disorder, and schizophrenia. Studies with identical twins raised in different environments indicate that individuals may inherit genetic potential or vulnerabilities, but environmental conditions may influence the expression of the genotype. Thus, phenotypic expression depends on a combination of nature and nuture.

■ ■

Review

■

What is incomplete dominance? codominance? What is the difference between multiple alleles and polygenic inheritance? What is pleiotropy? epistasis?

■

What is a norm of reaction?

■ ■

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

11.1 (page 238) 1 Define the terms phenotype, genotype, locus, allele, dominant allele, recessive allele, homozygous, and heterozygous. ■ Genes are in chromosomes; the site a gene occupies in the chromosome is its locus. Different forms of a particular gene are alleles; they occupy corresponding loci on homologous chromosomes. ■ An individual that carries two identical alleles is homozygous for that locus. If the two alleles are different, that individual is heterozygous for that locus. One allele, the dominant allele, may mask the expression of the ■ other allele, the recessive allele, in a heterozygous individual. For this reason two individuals with the same appearance, or phenotype, may differ from each other in their genetic makeup (that is, their combination of alleles), or genotype. 2 Describe Mendel’s principles of segregation and independent assortment. ■ According to Mendel’s principle of segregation, during meiosis the alleles for each locus separate, or segregate, from each other. When haploid gametes are formed, each contains only one allele for each locus. ■ According to Mendel’s principle of independent assortment, alleles of different loci are distributed randomly into gametes. This can result in genetic recombination, the production of new allele combinations that were not present in the parental (P) generation. Learn more about independent assortment by clicking on the figure in CengageNOW.

3 Distinguish among monohybrid, dihybrid, and test crosses. ■

■

A cross between homozygous parents (P generation) that differ from each other with respect to their alleles at one locus is called a monohybrid cross; if they differ at two loci, it is called a dihybrid cross. The first generation of offspring, called the F1 generation, is heterozygous; the generation produced by a cross of two F1 individuals is the F2 generation. A test cross between an individual of unknown genotype and a recessive individual helps determine the unknown genotype. Learn more about monohybrid, dihybrid, and test crosses by clicking on the figures in CengageNOW.

4 Explain Mendel’s principles of segregation and independent assortment, given what scientists now know about genes and chromosomes. Segregation of alleles is a direct result of homologous chromosomes separating during meiosis. ■ Independent assortment occurs because there are two ways in which two pairs of homologous chromosomes can be arranged at metaphase I of meiosis. The orientation of homologous chromosomes on the metaphase plate determines the way chromosomes are distributed into haploid cells. ■

11.2 (page 247) 5 Apply the product rule and sum rule appropriately when predicting the outcomes of genetic crosses. ■ According to the product rule, the probability of two independent events occurring together can be calculated by multiplying the probabilities of each event occurring separately. ■ According to the sum rule, the probability of an outcome that can be obtained in more than one way can be calculated by adding the separate probabilities.

11.3 (page 249) 6 Define linkage and relate it to specific events in meiosis. Linkage is the tendency for a group of genes on the same chromosome to be inherited together. Independent assortment does not apply if two loci are linked close together on the same pair of homologous chromosomes. ■ Recombination of linked genes can result from crossing-over (breaking and rejoining of homologous chromatids) in meiotic prophase I. (Recall from Section 11.1 that recombination can also result from independent assortment of unlinked genes.) 7 Show how data from a test cross involving alleles of two loci can be used to distinguish between independent assortment and linkage. ■ To distinguish between independent assortment of unlinked genes and linked genes, perform a two-point test cross between an individual that is heterozygous at both loci and an individual that is homozygous recessive for both. ■ Linkage is recognized when an excess of parental-type offspring and a deficiency of recombinant-type offspring are produced in a two-point test cross. ■

8 Discuss the genetic determination of sex and the inheritance of Xlinked genes in mammals. ■ The sex of humans and other mammals is determined by the X and Y chromosomes. Normal female mammals have two X chromosomes; normal males have one X and one Y. The fertilization of an X-bearing egg by an X-bearing sperm results in a female (XX) zygote. The fertilization of an X-bearing egg by a Y-bearing sperm results in a male (XY) zygote. ■ The Y chromosome determines male sex in mammals. The X chromosome contains many important genes unrelated to sex determination that are required by both males and females. A male receives all his X-linked genes from his mother. A female receives X-linked genes from both parents.

11.4 (page 255) 9 Explain some of the ways genes may interact to affect the phenotype; discuss how a single gene can affect many features of the organism simultaneously. ■ Pleiotropy is the ability of one gene to have several effects on different characters. Most cases of pleiotropy can be traced to a single cause, such as a defective enzyme. Alternatively, alleles of many loci may interact to affect the phenotypic expression of a single character. 10 Distinguish among incomplete dominance, codominance, multiple alleles, epistasis, and polygenic inheritance. ■ Dominance does not always apply; some alleles demonstrate incomplete dominance, in which the heterozygote is intermediate in phenotype, or codominance, in which the heterozygote simultaneously expresses the phenotypes of both homozygotes. ■ Multiple alleles, three or more alleles that can potentially occupy a particular locus, may exist in a population. A diploid individual has any two of the alleles; a haploid individual or gamete has only one.

In epistasis, an allele of one locus can mask the expression of alleles of a different locus. ■ In polygenic inheritance, multiple independent pairs of genes may have similar and additive effects on the phenotype. 11 Describe norm of reaction and give an example. ■ The range of phenotypic possibilities that can develop from a single genotype under different environmental conditions is known as the norm of reaction. ■ Many genes are involved in the inheritance of height in humans. Also, height is modified by a variety of environmental conditions, such as diet and general health. The genes that affect height set the norm of reaction—that is, the limits—for the phenotype, and the environment molds the phenotype within its norm of reaction. ■

T E S T YO U R U N D E R S TA N D I N G 1. One of the autosomal loci controlling eye color in fruit flies has two alleles, one for brown eyes and the other for red eyes. Fruit flies from a true-breeding line with brown eyes were crossed with flies from a true-breeding line with red eyes. The F1 flies had red eyes. What conclusion can be drawn from this experiment? (a) these alleles underwent independent assortment (b) these alleles underwent segregation (c) these genes are X-linked (d) the allele for red eyes is dominant to the allele for brown eyes (e) all of the preceding are true 2. The F1 flies described in question 1 were mated with browneyed flies from a true-breeding line. What phenotypes would you expect the offspring to have? (a) all red eyes (b) all brown eyes (c) half red eyes and half brown eyes (d) red-eyed females and brown-eyed males (e) brown-eyed females and red-eyed males 3. The type of cross described in question 2 is (a) an F2 cross (b) a dihybrid cross (c) a test cross (d) a two-point test cross (e) none of these 4. Individuals of genotype AaBb were crossed with aabb individuals. Approximately equal numbers of the following classes of offspring were produced: AaBb, Aabb, aaBb, and aabb. These results illustrate Mendel’s principle(s) of (a) linkage (b) independent assortment (c) segregation (d) a and c (e) b and c 5. Assume that the ratio of females to males is 1:1. A couple already has two daughters and no sons. If they plan to have a total of six children, what is the probability they will have four more girls? 1 1 1 (a) –14 (b) –18 (c) –– 16 (d) –– 32 (e) –– 64 6. Red–green color blindness is an X-linked recessive disorder in humans. Your friend is the daughter of a color-blind father. Her mother had normal color vision, but her maternal grandfather was color-blind. What is the probability your friend is color-blind? (a) 1 (b) –12 (c) –14 (d) –43 (e) 0

7. When two long-winged flies were mated, the offspring included 77 with long wings and 24 with short wings. Is the short-winged condition dominant or recessive? What are the genotypes of the parents? 8. The long hair of Persian cats is recessive to the short hair of Siamese cats, but the black coat color of Persians is dominant to the brown-and-tan coat color of Siamese. Make up appropriate symbols for the alleles of these two unlinked loci. If a pure black, long-haired Persian is mated to a pure brown-and-tan, short-haired Siamese, what will be the appearance of the F1 offspring? If two of these F1 cats are mated, what is the chance that a long-haired, brown-and-tan cat will be produced in the F2 generation? (Use the shortcut probability method to obtain your answer; then, check it with a Punnett square.) 9. Mr. and Mrs. Smith are concerned because their own blood types are A and B, respectively, but their new son, Richard, is blood type O. Could Richard be the child of these parents? 10. A walnut comb rooster is mated to three hens. Hen A, which has a walnut comb, has offspring in the ratio of 3 walnut to 1 rose. Hen B, which has a pea comb, has offspring in the ratio of 3 walnut to 3 pea to 1 rose to 1 single. Hen C, which has a walnut comb, has only walnut comb offspring. What are the genotypes of the rooster and the three hens? 11. Individuals of genotype AaBb were mated to individuals of genotype aabb. One thousand offspring were counted, with the following results: 474 Aabb, 480 aaBb, 20 AaBb, and 26 aabb. What type of cross is this? Are these loci linked? What are the two parental classes and the two recombinant classes of offspring? What is the percentage of recombination between these two loci? How many map units apart are they? 12. Genes A and B are 6 map units apart, and A and C are 4 map units apart. Which gene is in the middle if B and C are 10 map units apart? Which is in the middle if B and C are 2 map units apart?

CRITICAL THINKING 1. Sketch a series of diagrams showing each of the following, making sure to end each series with haploid gametes: (a) How a pair of alleles for a single locus segregate in meiosis (b)How the alleles of two unlinked loci assort independently in meiosis (c) How the alleles of two linked loci undergo genetic recombination 2. Can you always ascertain an organism’s genotype for a particular locus if you know its phenotype? Conversely, if you are given an organism’s genotype for a locus, can you always reliably predict its phenotype? Explain. 3. Compare the mechanisms of genetic recombination in linked and unlinked genes. 4. EVOLUTION LINK. Darwin’s theory of evolution by natural selection is based on four observations about the natural world. One of these is that each individual has a combination of traits that makes it uniquely different. Darwin recognized that much of this variation among individuals must be inherited, but he did not know about Mendel’s mechanism of inheritance. Based on what you have learned in this chapter, briefly explain the variation among individuals that Darwin observed. 5. ANALYZING DATA. Using the graph in Figure 11-20, determine how many offspring were involved in the hypothetical cross studying skin color. What percentage had the lightest skin possible? the darkest skin? 6. ANALYZING DATA. The following figure shows a cross between a homozygous black and a homozygous yellow Labrador retriever. What is the phenotype of the F1 generation? What are the phenotypes and the phenotypic ratio of the F2 generation? Inheritance

of coat color in Labrador retrievers is an example of what type of inheritance pattern? Homozygous parents: BBEE X bbee

F1 puppies: BbEe

Allele combinations possible among F2 puppies: BE

Be

bE

be

BE

BBEE

BBEe

BbEE

BbEe

Be

BBEe

BBee

BbEe

Bbee

bE

BbEE

BbEe

bbEE

bbEe

be

BbEe

Bbee

bbEe

bbee

Additional questions are available in CengageNOW at www.cengage.com/ login.

12

DNA: The Carrier of Genetic Information

Dr. Gopal Murti/Visuals Unlimited

Electron micrograph of DNA replication. During replication, two DNA molecules are synthesized from the original parent molecule. Replication is occurring at the Y-shaped structure, which is called a replication fork.

KEY CONCEPTS

A

fter the rediscovery of Mendel’s principles in 1900, geneticists conducted experiments to learn how genes are arranged in chro-

12.1 Starting in the 1920s, evidence began to accumulate that DNA is the hereditary material.

mosomes and how they are transmitted from generation to generation.

12.2 The DNA molecule consists of two strands that wrap

half of the 20th century: What are genes made of? How do genes work?

around each other to form a double helix; the order of its building blocks provides for the storage of genetic information. The DNA building blocks consist of four different nucleotide subunits, designated T, C, A, and G. The pairing of nucleotide subunits occurs based on precise pairing rules: T pairs with A, and C pairs with G.

The studies of inheritance patterns described in Chapter 11 did not answer

12.3 DNA replication, which results in two identical

tion in a form that the cell could retrieve and use. But scientists also had

double-stranded DNA molecules, provides the molecular mechanism for passing genetic information from one generation to the next.

However, basic questions remained unanswered through most of the first

these questions. However, they provided a foundation of knowledge that enabled scientists to make predictions about the molecular (chemical) nature of genes and how genes function. Scientists generally agreed that genes would have to store informato account for other properties of genes. For example, experiments on a variety of organisms had demonstrated that genes are usually stable and passed unchanged from generation to generation. However, occasionally a gene converted to a different form; such genetic changes, called mutations, were then transmitted unchanged to future generations.

As the science of genetics was developing, biochemists were

thought unremarkable, is the molecular basis of inheritance. We

making a growing effort to correlate the known properties of

explore the unique features of DNA, including its structure and

genes with the nature of various biological molecules. What kind of

replication (see photograph), that enable it to carry out this role.

molecule could store information? How could that information be retrieved and used to direct cell functions? What kind of molecule could be relatively stable but have the capacity to change, resulting in a mutation, under certain conditions? As they learned more about the central role of proteins in

12.1 EVIDENCE OF DNA AS THE HEREDITARY MATERIAL

virtually every aspect of cell structure and metabolism, some sci-

■ ■ LEARNING OBJECTIVES

entists considered proteins the prime candidates for the genetic

1

material. However, protein did not turn out to be the molecule that governs inheritance. In this chapter we discuss how researchers discovered that deoxyribonucleic acid (DNA), a nucleic acid once

2

Summarize the evidence that accumulated during the 1940s and early 1950s demonstrating that DNA is the genetic material. State the questions that these classic experiments addressed: Griffith’s transformation experiment, Avery’s contribution to Griffith’s work, and the Hershey–Chase experiments.

KEY EXPERIMENTS QUESTION: Can a genetic trait be transmitted from one bacterial strain to another? HYPOTHESIS: The ability of pneumococcus bacteria to cause disease can be transmitted from the virulent strain (smooth, or S cells) to the avirulent strain (rough, or R cells). EXPERIMENT: Griffith performed four experiments on mice, using the two strains of pneumococci: (1) injection of mice with live rough cells, (2) injection with live smooth cells, (3) injection with heat-killed smooth cells, and (4) injection with both live rough cells and heat-killed smooth cells. Experiment 1

Experiment 2

Experiment 3

Experiment 4

R cells injected

S cells injected

Heat-killed S cells injected

R cells and heat-killed S cells injected

Mouse lives

Mouse dies

Mouse lives

Mouse dies

RESULTS AND CONCLUSION: Although neither the rough strain nor the heat-killed smooth strain could kill a mouse, a combination of the two did. Autopsy of the dead mouse showed the presence of living S-strain pneumococci. These results indicated that some substance in the heat-killed S cells had transformed the living R cells into a virulent form.

FIGURE 12-1 Animated Griffith’s transformation experiments Griffith was trying to develop a vaccine against pneumonia when he serendipitously discovered the phenomenon of transformation.

During the 1930s and early 1940s, most geneticists paid little attention to DNA, convinced that the genetic material must be protein. Given the accumulating evidence that genes control production of proteins (discussed in Chapter 13), it certainly seemed likely that genes themselves must also be proteins. Scientists knew proteins consisted of more than 20 different kinds of amino acids in many different combinations, which conferred unique properties on each type of protein. Given their complexity and diversity compared with other molecules, proteins seemed to be the “stuff ” of which genes are made. In contrast, scientists had established that DNA and other nucleic acids were made of only four nucleotides, and what was known about their arrangement made them relatively uninteresting to most researchers. For this reason, several early clues to the role of DNA were not widely noticed.

DNA is the transforming principle in bacteria One of these clues had its origin in 1928, when Frederick Griffith, a British medical officer, made a curious observation concerning two strains of pneumococcus bacteria (FIG. 12-1). A smooth (S) strain, named for its formation of smooth colonies on a solid growth medium, exhibited virulence,

© Eye of Science/Photo Researchers, Inc.

the ability to cause disease, and often death, in its host. When living cells of this strain were injected into mice, the animals contracted pneumonia and died. Not surprisingly, the injected animals survived if the cells were first killed with heat. A related rough (R) strain of bacteria, which forms colonies with a Bacteriophage rough surface, exhibited avirulence, or inability to produce pathogenic effects; mice injected with either living or heat-killed cells of this strain survived. Phage genetic However, when Griffith injected mice with a mixmaterial ture of heat-killed virulent S cells and live avirulent R cells, a high proportion of the mice died. Griffith then isolated living S cells from the dead mice. Because neither the heat-killed S strain nor the living R strain could be converted to the living virulent form when injected by itself in the control Bacterial cell (E. coli ) experiments, it seemed that something in the heatkilled cells converted the avirulent cells to the lethal form. This type of permanent genetic change in which the properties of one strain of dead cells are conferred on a different strain of living cells is called transformation. Scientists hypothesized that some chemical substance (a transforming principle, or factor) was transferred from the dead bacteria to the living cells and caused transformation. In 1944, Oswald T. Avery and his colleagues Colin M. MacLeod and Maclyn McCarty chemically identified Griffith’s transforming factor as DNA. They did this through a series of careful experiments in which they lysed (split open) S cells and separated the cell contents into several fractions: lipids, proteins, polysaccharides, and nucleic acids (DNA and RNA). They tested each fraction to see if it could transform living R cells into S cells. The experiments using lipids, polysaccharides, and proteins did not cause transformation. However, when Avery treated living R cells with nucleic acids extracted from S cells, the R cells were transformed into S cells. Although today scientists consider these results to be the first definitive demonstration that DNA is the genetic material, not all scientists of the time were convinced. Many thought the findings might apply only to bacteria and might not have any relevance for the genetics of eukaryotes. During the next few years, new evidence accumulated that the haploid nuclei of pollen grains and gametes such as sperm contain only half the amount of DNA found in diploid somatic cells of the same species. (Somatic cells are body cells and never become gametes.) Because scientists generally accepted that genes are located on chromosomes, these findings correlating DNA content with chromosome number provided strong circumstantial evidence for DNA’s importance in eukaryotic inheritance.

0.5 μm

FIGURE 12-2 Bacteriophages on a bacterial cell This colorized TEM shows several phages attached to the bacterium Escherichia coli. Note the viral genetic material being injected into the bacterium.

As shown in FIGURE 12-3, they labeled the viral protein of one sample of phages with 35S, a radioactive isotope of sulfur, and the viral DNA of a second sample with 32P, a radioactive isotope of phosphorus. Recall from Chapter 3 that proteins contain sulfur as part of the amino acids cysteine and methionine and that nucleic acids contain phosphate groups. The phages in each sample attached to bacteria, and the researchers shook the phages off by agitating the sample in a blender. Then they centrifuged the samples. In the sample in which they had labeled the proteins with 35S, they subsequently found radioactivity in the supernatant, indicating that the protein had not entered the cells. In the sample in which they had labeled the DNA with 32P, they found radioactivity associated with the bacterial cells (in the pellet): DNA had actually entered the cells. Hershey and Chase concluded that phages inject their DNA into bacterial cells, leaving most of their protein on the outside. This finding emphasized the significance of DNA in viral reproduction, and many scientists saw it as an important demonstration of DNA’s role as the hereditary material.

Review ■

■

DNA is the genetic material in certain viruses In 1952, geneticists Alfred Hershey and Martha Chase performed a series of elegant experiments on the reproduction of viruses that infect bacteria, known as bacteriophages or phages (discussed in Chapter 24). When they planned their experiments, they knew that phages reproduce inside a bacterial cell, eventually causing the cell to break open and release large numbers of new viruses. Because electron microscopic studies had shown that only part of an infecting phage actually enters the cell, they reasoned that the genetic material should be included in that portion (FIG. 12-2).

How did the experiments of Avery and his colleagues point to DNA as the essential genetic material? How did the Hershey–Chase experiments establish that DNA is the genetic material in bacteriophages?

12.2 THE STRUCTURE OF DNA ■ ■ LEARNING OBJECTIVES 3 Explain how nucleotide subunits link to form a single DNA strand. 4 Describe how the two strands of DNA are oriented with respect to each 5

other. State the base-pairing rules for DNA and describe how complementary bases bind to each other.

KEY EXPERIMENT QUESTION: Is DNA or protein the genetic material in bacterial viruses (phages)? HYPOTHESIS: DNA is the genetic material in bacterial viruses. EXPERIMENT: Hershey and Chase produced phage populations with either radioactively labeled DNA or radioactively labeled protein. In both cases, they infected bacteria with the phages and then determined whether DNA or protein was injected into bacterial cells to direct the formation of new viral particles.

1 Radioactively labeled phages infect bacterial cells.

2 Cells agitated in blender and centrifuged. Bacteria, which are heavier, settle into pellet. Phages and phage parts, which are lighter, remain in supernatant.

35S

Phages from pellet inject bacteria into genetic material.

35S

Experiment A, phages with 35S-labeled protein coat (green)

35S-labeled

protein in supernatant

32P

Experiment B, phages with 32P-labeled DNA (red)

3 Pellets and supernatants tested for radioactivity.

Bacteria in pellet do not contain 35S label.

No radioactively labeled protein enters cell.

Phages from pellet inject genetic material. 32P

Bacteria in pellet contain 32P-labeled DNA.

Some newly assembled viruses have radioactively labeled DNA.

RESULTS AND CONCLUSION: The researchers could separate phage protein coats labeled with the radioactive isotope 35S from infected bacterial cells without affecting viral reproduction. However, they could not separate viral DNA labeled with the radioactive isotope 32P from infected bacterial cells. This demonstrated that viral DNA enters the bacterial cells and is required for the synthesis of new viral particles. Thus, DNA is the genetic material in phages.

FIGURE 12-3 Animated The Hershey–Chase experiments

Scientists did not generally accept DNA as the genetic material until 1953, when American scientist James Watson and British scientist Francis Crick, both working in England, proposed a model for its structure that had extraordinary explanatory power. The story of how the structure of DNA was figured out is one of the most remarkable chapters in the history of modern biology (TABLE 12-1).

As you will see in the following discussion, scientists already knew a great deal about DNA’s physical and chemical properties when Watson and Crick became interested in the problem; in fact, they did not conduct any experiments or gather any new data. Their allimportant contribution was to integrate all the available information into a model that demonstrated how the molecule can both

TABLE 12-1

Date

A Time Line of Selected Historical DNA Discoveries

1928 1944 1949 1952 1952 1953

1958 1962

1969

Friedrich Miescher reports discovery of new substance, nuclein, from cell nuclei. Nuclein is now known to be a mixture of DNA, RNA, and proteins. Frederick Griffith finds a substance in heat-killed bacteria that “transforms” living bacteria. Oswald Avery, Colin MacLeod, and Maclyn McCarty chemically identify Griffith’s transforming principle as DNA. Erwin Chargaff reports relationships among DNA bases that provide a clue to the structure of DNA. Alfred Hershey and Martha Chase demonstrate that DNA, not protein, is involved in viral reproduction. Rosalind Franklin produces X-ray diffraction images of DNA. James Watson and Francis Crick propose a model of the structure of DNA; this contribution is widely considered the start of a revolution in molecular biology that continues to the present. Matthew Meselson and Franklin Stahl demonstrate that DNA replication is semiconservative. James Watson, Francis Crick, and Maurice Wilkins are awarded the Nobel Prize in Medicine for discoveries about the molecular structure of nucleic acids.* Alfred Hershey is awarded the Nobel Prize in Medicine for discovering the replication mechanism and genetic structure of viruses.

H3C O

Discovery O

1871

O

5′

H 5′ O CH 2

P

4′

O–

H

N

O Thymine

O

H 3′

H 2′

P

H

1′

H

H

N N

O CH2

O– H

N

O

H

H

O

P

H

N Adenine

H

H

H

H

O

CH2

O Phosphate group

H

N

H

O O

H

N

H

H

N

N

H

O–

Nucleotide

N O

H

H

O Cytosine O

H

N

H O Phosphodiester linkage

O

P

N O

O–

CH2

O

N

H

N N Guanine H

H

H H H 3′ Deoxyribose OH H (sugar) H

3′

*Rosalind Franklin could not share the prize because she was deceased.

FIGURE 12-4 The nucleotide subunits of DNA

carry information for making proteins and serve as its own template (pattern or guide) for its duplication.

Nucleotides can be covalently linked in any order to form long polymers As discussed in Chapter 3, each DNA building block is a nucleotide consisting of the pentose sugar deoxyribose, a phosphate, and one of four nitrogenous bases (FIG. 12-4). It is conventional to number the atoms in a molecule using a system devised by organic chemists. Accordingly, in nucleic acid chemistry the individual carbons in each sugar and each base are numbered. The carbons in a base are designated by numerals, but the carbons in a sugar are distinguished from those in the base by prime symbols, such as 2¿. The nitrogenous base is attached to the 1¿ carbon of the sugar, and the phosphate is attached to the 5¿ carbon. The bases include two purines, adenine (A) and guanine (G), and two pyrimidines, thymine (T) and cytosine (C). The nucleotides are linked by covalent bonds to form an alternating sugar–phosphate backbone. The 3¿ carbon of one sugar is bonded to the 5¿ phosphate of the adjacent sugar to form a 3¿, 5¿ phosphodiester linkage. The result is a polymer of indefinite length, with the nucleotides linked in any order. Scientists now know that most DNA molecules found in cells are millions of bases long. Figure 12-4 also shows that a single polynucleotide chain is directional. No matter how long the chain may be, one end, the 5¿ end, has a 5¿ carbon attached to a phosphate and the other, the 3¿ end, has a 3¿ carbon attached to a hydroxyl group.

A single strand of DNA consists of a backbone (superimposed on blue screen) of phosphate groups alternating with the sugar deoxyribose (green). Phosphodiester linkages (pink) join sugars of adjacent nucleotides. Linked to the 1¿ carbon of each sugar is one of four nitrogenous bases (top to bottom): thymine, adenine, cytosine, and guanine. (The nucleotide containing the base adenine is highlighted yellow.) Note the polarity of the polynucleotide chain, with the 5¿ end at the top of the figure and the 3¿ end at the bottom.

In 1949, Erwin Chargaff and his colleagues at Columbia University had determined the base composition of DNA from several organisms and tissues. They found a simple relationship among the bases that turned out to be an important clue to the structure of DNA. Regardless of the source of the DNA, in Chargaff ’s words the “ratios of purines to pyrimidines and also of adenine to thymine and of guanine to cytosine were not far from 1” (TABLE 12-2).

TABLE 12-2

Base Compositions in DNA from Selected Organisms Percentage of DNA Bases

Source of DNA E. coli Yeast Sea urchin sperm Herring sperm Human liver Corn (Zea mays)

A 26.1 31.3 32.5 27.8 30.3 25.6

T 23.9 32.9 31.8 27.5 30.3 25.3

G 24.9 18.7 17.5 22.2 19.5 24.5

C 25.1 17.1 18.2 22.6 19.9 24.6

Ratios A/T 1.09 0.95 1.02 1.01 1.00 1.01

G/C 0.99 1.09 0.96 0.98 0.98 1.00

In other words, in double-stranded DNA molecules, the number of purines equals the number of pyrimidines, the number of adenines equals the number of thymines (A equals T), and the number of guanines equals the number of cytosines (G equals C). These equalities are called Chargaff ’s rules.

RESEARCH METHOD Why Is It Used?

DNA is made of two polynucleotide chains intertwined to form a double helix Key information about the structure of DNA came from X-ray diffraction studies on crystals of purified DNA, carried out by British scientist Rosalind Franklin at King’s College from 1951 to 1953 (FIG. 12-5). X-ray diffraction, a powerful method for determining the 3-D structure of a molecule, can determine the distances between the atoms of molecules arranged in a regular, repeating crystalline structure (FIG. 12-6). X-rays have such short wavelengths that they can be scattered by the electrons surrounding the atoms in a molecule. Atoms with dense electron clouds (such as phosphorus and oxygen) tend to deflect electrons more strongly than atoms with lower atomic numbers. Exposing a crystal to an intense beam of X-rays causes the regular arrangement of its atoms to diffract, or scatter,

How Is It Done?

X-ray diffraction can be used to determine the regular array of atoms in a crystalline sample of, for example, DNA. Because each type of crystal has its own characteristic pattern, the three-dimensional structure of the molecule being studied can be deduced. 1 Researchers direct a narrow beam of ●

X-rays at a single crystal of DNA. X-rays are diffracted (bent) at specific angles based on the density of electrons of different atoms. Important clues about DNA structure are provided by detailed mathematical analysis of measurements of the spots, which are formed by X-rays hitting the photographic plate

Beam of X-rays

X-ray source

DNA sample

X-rays diffracted by DNA Photographic plate shows atomic array of DNA.

2 X-ray diffraction image of DNA. This ●

© Science Source/Photo Researchers, Inc.

Dr. S. Dover, Division of Biomolecular Sciences, Kings College, London

diagonal pattern of spots stretching from 11 to 5 and from 1 to 7 (as on a clock face) provides evidence for the helical structure of DNA. The elongated horizontal patterns at the top and bottom indicate that the purine and pyrimidine bases are stacked 0.34 nm apart and are perpendicular to the axis of the DNA molecule.

FIGURE 12-5 Rosalind Franklin Franklin was a gifted scientist whose contributions helped elucidate the double helix structure of DNA.

FIGURE 12-6 How X-ray diffraction works

the X-rays in specific ways. The pattern of diffracted X-rays appears on photographic film as dark spots. Mathematical analysis of the pattern and distances between the spots yields the precise distances between atoms and their orientation within the molecules. Franklin had already produced X-ray crystallographic films of DNA patterns when Watson and Crick began to pursue the problem of DNA structure. Her pictures clearly showed that DNA has a type of helical structure, and three major types of regular, repeating patterns in the molecule (with the dimensions 0.34 nm, 3.4 nm, and 2.0 nm) were evident. Franklin and colleague Maurice Wilkins had inferred from these patterns that the nucleotide bases (which are flat molecules) are stacked like the rungs of a ladder. Using this information, Watson and Crick began to build scale models of the DNA components and then fit them together to correlate with the experimental data (FIG. 12-7). After several trials, they worked out a model that fit the existing data (FIG. 12-8). The nucleotide chains conformed to the dimensions of the X-ray data only if each DNA molecule consisted of two polynucleotide chains arranged in a coiled double helix. In their model, the sugar–phosphate backbones of the two chains form the outside of the helix. The bases belonging to the two chains associate as pairs along the central axis of the helix. The reasons for the repeating patterns of 0.34 nm and 3.4 nm measurements are readily apparent from the model: each pair of bases is exactly 0.34 nm from the adjacent pairs above and below. Because exactly 10 base pairs are present in each full turn of the helix, each turn constitutes 3.4 nm of length. To fit the data, the two chains must run in opposite directions; therefore, each end of the double helix must have an exposed 5¿ phosphate on one strand and an exposed 3¿ hydroxyl

group (¬OH) on the other. Because the two strands run in opposite directions, they are antiparallel to each other (FIG. 12-9a).

In double-stranded DNA, hydrogen bonds form between A and T and between G and C Other features of the Watson and Crick model integrated critical information about the chemical composition of DNA with the Sugar– phosphate backbone

Minor groove

Major groove

3.4 nm

© A. Barrington Brown/Photo Researchers, Inc.

0.34 nm

FIGURE 12-7 James Watson and Francis Crick Watson (left) and Crick (right) are shown with their model of DNA’s double helix.

2.0 nm = hydrogen

= atoms in base pairs

= oxygen

= carbon

= phosphorus

FIGURE 12-8 Animated A three-dimensional model of the DNA double helix The measurements match those derived from X-ray diffraction images.

KEY POINT

Base pairing and the sequence of bases in DNA provide a foundation for understanding both DNA replication and the inheritance of genetic material.

G H H

A

O CH2

P

O –O P O O CH2

O

O C

H H H

O H H

A

T

O –O P O O CH2

H H H

G

T

H H

C

3′

G

Adenine

H

CH2 O P O–

C

C N

N

N C

O O H

N N

O

H

C

C

O

C

(a) The two sugar–phosphate chains run in opposite directions. This orientation permits the complementary bases to pair.

C

H

C

H H

N

O Deoxyribose

Guanine

H

C N

O

Cytosine H O

N C C

H

C C

H

C

N

C

H

H

N C

N N

H

C

N HO H Deoxyribose

OH

O

HO H Deoxyribose

CH2 O O P O– O

5′

H

C

N

C H

CH2 O O P O– O

A

H H H

Thymine H

H

CH2 O O P O– O

O P O O CH2

O

CH2 O P O–

G

O P O O CH2

–O

T

O

–O

–O

H H H H C H

N

H H

OH

O O Deoxyribose

(b) Hydrogen bonding between base pairs adenine (A) and thymine (T) (top) and guanine (G) and cytosine (C) (bottom). The AT pair has two hydrogen bonds; the GC pair has three.

FIGURE 12-9 Animated Base pairing and hydrogen bonding The two strands of a DNA double helix are hydrogen-bonded between the bases.

X-ray diffraction data. The X-ray diffraction studies indicated that the double helix has a precise and constant width, as shown by the 2.0 nm measurements. This finding is actually consistent with Chargaff ’s rules. As Figure 12-4 shows, each pyrimidine (cytosine or thymine) contains only one ring of atoms, whereas each purine

(guanine or adenine) contains two rings. Study of the models made it clear to Watson and Crick that if each rung of the ladder contained one purine and one pyrimidine, the width of the helix at each base pair would be exactly 2.0 nm. By contrast, the combination of two purines (each of which is 1.2 nm wide) would be wider

than 2.0 nm and that of two pyrimidines would be narrower, so the diameter would not be constant. Further examination of the model showed that adenine can pair with thymine (and guanine with cytosine) in such a way that hydrogen bonds form between them; the opposite combinations, cytosine with adenine and guanine with thymine, do not lead to favorable hydrogen bonding. The nature of the hydrogen bonding between adenine and thymine and between guanine and cytosine is shown in FIGURE 12-9b. Two hydrogen bonds form between adenine and thymine, and three between guanine and cytosine. This concept of specific base pairing neatly explains Chargaff ’s rules. The amount of cytosine must equal the amount of guanine because every cytosine in one chain must have a paired guanine in the other chain. Similarly, every adenine in the first chain must have a thymine in the second chain. The sequences of bases in the two chains exhibit complementary base pairing—that is, the sequence of nucleotides in one chain dictates the complementary sequence of nucleotides in the other. For example, if one strand has this sequence, 3¿¬AGCTAC¬5¿

then the other strand has the complementary sequence: 5¿¬TCGATG¬3¿

The double-helix model strongly suggested that the sequence of bases in DNA provides for the storage of genetic information and that this sequence ultimately relates to the sequences of amino acids in proteins. Although restrictions limit how the bases on opposite strands pair with each other, the number of possible linear sequences of bases in a strand is virtually unlimited. Because a DNA molecule in a cell can be millions of nucleotides long, it can store enormous amounts of information, usually consisting of hundreds of genes.

Review What types of subunits make up a single strand of DNA? How are the subunits linked? What is the structure of double-stranded DNA, as determined by Watson and Crick? How do Chargaff’s rules relate to the structure of DNA?

■

■

■

12.3 DNA REPLICATION ■ ■ LEARNING OBJECTIVES 6 Cite evidence from Meselson and Stahl’s experiment that enabled scien-

7 8 9 10

tists to differentiate between semiconservative replication of DNA and alternative models. Summarize how DNA replicates and identify some unique features of the process. Explain the complexities of DNA replication that make it (a) bidirectional and (b) continuous in one strand and discontinuous in the other. Discuss how enzymes proofread and repair errors in DNA. Define telomere, and describe the possible connections between telomerase and cell aging and between telomerase and cancer.

Two immediately apparent and distinctive features of the Watson– Crick model made it seem plausible that DNA is the genetic mate-

rial. We have already mentioned that the sequence of bases in DNA can carry coded information. The model also suggested a way in which the sequence of nucleotides in DNA could be precisely copied, a process known as DNA replication. The connection between DNA replication and the behavior of chromosomes in mitosis was obvious to Watson and Crick. A chromosome becomes duplicated so that it consists of two identical sister chromatids that later separate at anaphase; the genetic material must be precisely duplicated and distributed to the daughter cells. They noted, in a classic and now famous understatement at the end of their first brief paper, “It has not escaped our notice that the specific pairing we have postulated immediately suggests a possible copying mechanism for the genetic material.” The model suggested that because the nucleotides pair with each other in complementary fashion, each strand of the DNA molecule could serve as a template for synthesizing the opposite strand. It would simply be necessary for the hydrogen bonds between the two strands to break (recall from Chapter 2 that hydrogen bonds are relatively weak) and the two chains to separate. Each strand of the double helix could then pair with new complementary nucleotides to replace its missing partner. The result would be two DNA double helices, each identical to the original one and consisting of one original strand from the parent molecule and one newly synthesized complementary strand. This type of information copying is called semiconservative replication (FIG. 12-10a).

Meselson and Stahl verified the mechanism of semiconservative replication Although the semiconservative replication mechanism suggested by Watson and Crick was (and is) a simple and compelling model, experimental proof was needed to establish that DNA in fact replicates in that manner. Researchers first needed to rule out other possibilities. With conservative replication, both parent (or old) strands would remain together, and the two newly synthesized strands would form a second double helix (FIG. 12-10b). As a third hypothesis, the parental and newly synthesized strands might become randomly mixed during the replication process—that is, dispersive replication (FIG. 12-10c). To discriminate among semiconservative replication and the other models, investigators had to distinguish between old and newly synthesized strands of DNA. One technique is to use a heavy isotope of nitrogen, 15N (ordinary nitrogen is 14N), to label the bases of the DNA strands, making them more dense. Using density gradient centrifugation, scientists can separate large molecules such as DNA on the basis of differences in their density (see Fig. 4-5). When DNA is mixed with a solution containing cesium chloride (CsCl) and centrifuged at high speed, the solution forms a density gradient in the centrifuge tube, ranging from a region of lowest density at the top to one of highest density at the bottom. During centrifugation, the DNA molecules migrate to the region of the gradient identical to their own density. In 1958, Matthew Meselson and Franklin Stahl at the California Institute of Technology grew the bacterium Escherichia coli on a medium that contained 15N in the form of ammonium chloride (NH4Cl). The cells used the 15N to synthesize bases, which then became incorporated into DNA (FIG. 12-11). The resulting DNA molecules, which contained heavy nitrogen, were extracted from some of the cells. When the researchers subjected them to density

(a) Hypothesis 1: Semiconservative replication Parental DNA

First generation

Second generation

KEY EXPERIMENT QUESTION: What is the mechanism of DNA replication? HYPOTHESIS: Figure 12-10 depicts three hypotheses of DNA replication and predicts the arrangement of old and newly synthesized DNA strands after one or two generations according to each hypothesis.

(b) Hypothesis 2: Conservative replication Parental DNA

First generation

EXPERIMENT: Meselson and Stahl grew bacteria (E. coli) in heavy nitrogen (15N) growth medium for many generations so that all the DNA strands would be heavy. Then they transferred some of the cells to light nitrogen (14N) medium so that the newly synthesized strands would be light. They isolated DNA from bacterial cells after 20 minutes (one generation) and 40 minutes (two generations) and centrifuged it to separate DNA into bands based on density.

Second generation

14N

(light) 14N –15N 15N (heavy) DNA hybrid DNA DNA

In density gradient centrifugation, the concentration of CsCl is highest at the bottom of the tube and lowest at the top. DNA molecules move to positions where their density equals that of the CsCl solution in which they are centrifuged.

(c) Hypothesis 3: Dispersive replication Parental DNA

First generation

Second generation

RESULTS AND CONCLUSION: Based on the observed density of the DNA molecules in each generation, Meselson and Stahl concluded that the semiconservative model accurately predicts the mechanism of DNA replication.

14 N (light) DNA 14

N –15N hybrid DNA

14

N –15N hybrid DNA

15

N (heavy) DNA

FIGURE 12-10 Alternative models of DNA replication The hypothesized arrangement of old (light blue) and newly synthesized (dark blue) DNA strands after one and two generations, according to (a) the semiconservative model, (b) the conservative model, and (c) the dispersive model.

gradient centrifugation, they accumulated in the high-density region of the gradient. The team transferred the rest of the bacteria (which also contained 15N-labeled DNA) to a different growth medium in which the NH4Cl contained the naturally abundant, lighter 14N isotope and then allowed them to undergo additional cell divisions. Meselson and Stahl expected the newly synthesized DNA strands to be less dense because they incorporated bases containing the lighter 14N isotope. Indeed, double-stranded DNA from cells isolated after one generation had an intermediate density, indicating they contained half as many 15N atoms as the “parent” DNA. This finding supported the semiconservative replication model, which predicted that each double helix would contain a previously synthesized strand (heavy in this case) and a newly synthesized strand (light in this case). It was also consistent with the disper-

Before transfer to 14N

One cell generation after transfer to 14N

Two cell generations after transfer to 14N

The location of DNA molecules within the centrifuge tube can be determined by UV optics. DNA solutions absorb strongly at 260 nm.

FIGURE 12-11 The Meselson–Stahl experiment

sive model, which would also yield one class of molecules, all with intermediate density. It was inconsistent with the conservative model, which predicted two classes of double-stranded molecules, those with two heavy strands and those with two light strands. After another cycle of cell division in the medium with the lighter 14N isotope, two types of DNA appeared in the density gradient, exactly as predicted by the semiconservative replication model. One consisted of DNA with a density intermediate between 15N-labeled DNA and 14N-labeled DNA, whereas the other contained only DNA with a density of 14N-labeled DNA. This finding refuted the dispersive model, which predicted that all strands would have an intermediate density.

Semiconservative replication explains the perpetuation of mutations The recognition that DNA could be copied by a semiconservative mechanism suggested how DNA could explain a third essential characteristic of genetic material—the ability to mutate. It was long known that mutations, or genetic changes, could arise in genes and then be transmitted faithfully to succeeding generations. For example, a mutation in the fruit fly (Drosophila melanogaster) produces vestigial wings. When the double-helix model was proposed, it seemed plausible that mutations could represent a change in the sequence of bases in the DNA. You could predict that if DNA is copied by a mechanism involving complementary base pairing, any change in the sequence of bases on one strand would produce a new sequence of complementary bases during the next replication cycle. The new base sequence would then transfer to daughter molecules by the same mechanism used to copy the original genetic material, as if no change had occurred. For the example in FIGURE 12-12, an adenine base in one of the DNA strands has been changed to guanine. This could occur by a rare error in DNA replication or by one of several other known mechanisms. As discussed later, certain enzymes correct errors when they occur, but not all errors are corrected properly. By one estimate, the rate of uncorrected errors that occur during DNA replication is equal to about one nucleotide in a billion. When the DNA molecule containing an error replicates (left side of Fig. 12-12), one of the strands gives rise to a molecule exactly like its parent strand; the other (mutated) strand gives rise to a molecule with a new combination of bases that is transmitted generation after generation.

DNA replication requires protein “machinery” Although semiconservative replication by base pairing appears simple and straightforward, the actual process is highly regulated and requires a “replication machine” containing many types of protein and enzyme molecules. Many essential features of DNA replication are common to all organisms, although prokaryotes and eukaryotes differ somewhat because their DNA is organized differently. In most bacterial cells, such as E. coli, most or all the DNA is in the form of a single, circular, double-stranded DNA molecule. In contrast, each unreplicated eukaryotic chromosome contains a single, linear, double-stranded molecule associated with at least as much protein (by mass) as DNA. We now present DNA replication, based on the current understanding of the process. Although scientists know much about DNA replication, many aspects of the process remain unclear. For example, in the unicellular yeast Saccharomyces cerevisiae, which is considered a relatively “simple” eukaryote, 88 genes are involved in DNA replication! Determining the roles and interactions of all those genes will require the efforts of many scientists over a long period. DNA strands must be unwound during replication DNA replication begins at specific sites on the DNA molecule, called origins of replication, where small sections of the double helix unwind. DNA helicases are helix-destabilizing enzymes

A

T

T

A

G

C

C

G

DNA replication in which mutation occurs A

T

T

G

Mutation

A

T

T

A

G

C

G

C

C

G

C

G

DNA replication in which mutation is propagated A

T

A

T

A

T

A

T

T

A

C

G

T

A

T

A

G

C

G

C

G

C

G

C

C

G

C

G

C

G

C

G

FIGURE 12-12 The perpetuation of a mutation The process of DNA replication can stabilize a mutation (bright yellow) so that it is transmitted to future generations.

(several have been identified) that bind to DNA at the origin of replication and break hydrogen bonds, thereby separating the two strands (TABLE 12-3). Both DNA strands replicate at the same time at the junction between the separated strands, which is a Y-shaped structure called the replication fork (see chapter opener photograph). Helicase travels along the helix, opening the double helix like a zipper during movement of the replication fork. Once DNA helicases separate the strands, single-strand binding (SSB) proteins bind to single DNA strands and stabilize them; this prevents the double helix from re-forming until the strands are replicated. SSB proteins also prevent the hydrolysis of the single-strand regions by other enzymes (nucleases; as we discuss later in the chapter, nucleases are involved in DNA repair). Watson and Crick recognized that in their double-helix model, the two DNA strands wrap around each other like the strands of a rope. If you try to pull the strands apart, the rope must either rotate or twist into tighter coils. You could expect similar results when complementary DNA strands are separated for replication. As the DNA strands unwind to open for replication, torsional strain from supercoiling (excessive twisting) occurs in another part of the DNA molecule. Enzymes called topoisomerases produce breaks in the DNA molecules and then rejoin the strands, relieving strain and effectively preventing supercoiling and knot formation during replication. There are two ways that topoisomerases reduce supercoiling. Some topoisomerases produce a temporary break in the polynucleotide backbone of a single strand of DNA, pass that strand through the excessively twisted part, and then reseal the break. Other topoisomerases break both DNA strands, pass some of the helix between the cut ends, and then reseal the break.

TABLE 12-3

Proteins Involved in DNA Replication 5′ 3′

Enzyme

Function

Helicase

Opens the double helix at replication forks by disrupting the hydrogen bonds that hold the two strands together. Binds to single strands of DNA and prevents the helix from re-forming before it can be used as a template for replication. Breaks one or both DNA strands, preventing excessive coiling during replication, and then rejoins them in a more relaxed configuration. Links nucleotide subunits to form a new DNA strand from a DNA template. Synthesizes short RNA primers on the lagging strand. Begins replication of the leading strand. Links Okazaki fragments by joining the 3¿ end of the new DNA fragment to the 5¿ end of the adjoining DNA.

Single-strand binding (SSB) protein Topoisomerase

DNA polymerase

3′ 5′

DNA primase 3′

DNA ligase

5′

DNA synthesis requires an RNA primer As mentioned, DNA polymerases add nucleotides only to the 3¿ end of an existing polynucleotide strand. Then, how is DNA synthesis initiated once the two strands are separated? The answer is that first a short piece of RNA (5 to 14 nucleotides) called an RNA primer is synthesized at the point where replication begins (FIG. 12-14). RNA, or ribonucleic acid (see Chapters 3 and 13), is a nucleic acid polymer consisting of nucleotide subunits that can associate by complementary base pairing with the single-strand DNA template. The RNA primer is synthesized by DNA primase, an enzyme that starts a new strand of RNA opposite a short stretch

Regardless of their modes of action, topoisomerases give replicating DNA a more relaxed configuration. DNA synthesis always proceeds in a 5¿ ¡ 3¿ direction The enzymes that catalyze the linking of successive nucleotide subunits are called DNA polymerases. They add nucleotides only to the 3¿ end of a growing polynucleotide strand, and this strand must be paired with the DNA template strand (FIG. 12-13). Nucleotides with three phosphate groups are substrates for the polymerization reaction. As the nucleotides become linked, two of the phosphates are removed. These reactions are strongly exergonic and do not 5′

3′

3′

3′ 5′

5′

A Base 3′ 5′

C

3′

need additional energy. Because the polynucleotide chain is elongated by the linkage of the 5¿ phosphate group of the next nucleotide subunit to the 3¿ hydroxyl group of the sugar at the end of the existing strand, the new strand of DNA always grows in the 5¿ ¡ 3¿ direction. Some DNA polymerases are very efficient in joining nucleotides to the growing polypeptide chain. DNA Pol III, which is one of five DNA polymerases that have been identified in the bacterium E. coli, can join 1200 nucleotides per minute.

G

5′ 3′ Nucleotide joined to growing 5′ chain by DNA polymerase

T

C

G

A

T

T

G

C

A

OH

Phosphates released

C 3′

3′

FIGURE 12-13 Animated A simplified view of DNA replication

C

G

C

OH

One nucleotide at a time is added to the 3¿ end of a growing chain.

A

T

5′

5′

KEY POINT

DNA replication requires several steps and involves several enzymes, proteins, and RNA primers.

DNA polymerase

Origin of replication on DNA molecule

3′

3′

5′

5′

Twist introduced into the helix by unwinding

Single-strand binding proteins

RNA primer

1

DNA synthesis begins at origin of replication.

2

Strands are separated at origin of replication and unwound by DNA helicase, which “walks” along DNA molecule preceding the DNA-synthesizing enzymes. Singlestranded regions are prevented from re-forming into double strands by singlestrand binding proteins, which bind to single DNA strands. Region of active DNA synthesis is associated with replication fork, formed at junction of single strands and double-stranded region. Both strands are synthesized in vicinity of fork 3′ direction). (in 5′

3

Completion of replication results in formation of two daughter molecules, each containing one old and one newly synthesized strand. Each double helix is a chromatid of a duplicated eukaryotic chromosome.

DNA polymerase 3′ 5′

3′

DNA helicase

3′ 5′

3′

RNA primer

Direction of replication

3′

3′

5′

5′

3′

3′

5′

5′

FIGURE 12-14 Animated An overview of DNA replication

of the DNA template strand. After a few nucleotides have been added, DNA polymerase displaces the primase and subsequently adds subunits to the 3¿ end of the short RNA primer. Specific enzymes later degrade the primer (discussed in the next section), and the space fills in with DNA. DNA replication is discontinuous in one strand and continuous in the other We mentioned earlier that the complementary DNA strands are antiparallel. DNA synthesis proceeds only in the 5¿ ¡ 3¿ direction, which means that the strand being copied is being read in the 3¿ ¡ 5¿ direction. Thus, it may seem necessary to copy one of the strands starting at one end of the double helix and the other strand starting at the opposite end. Some viruses replicate their DNA in this way, but this replication method is not workable in the extremely long DNA molecules in eukaryotic chromosomes. Instead, as previously mentioned, DNA replication begins at origins of replication, and both strands replicate at the same time at a replication fork (FIG. 12-15). The position of the replication fork is constantly moving as replication proceeds. Two identical DNA polymerase molecules catalyze replication. One of these adds nucleotides to the 3¿ end of the new strand that is always growing toward the replication fork. Because this strand is synthesized smoothly and continuously, it is called the leading strand. A separate DNA polymerase molecule adds nucleotides to the 3¿ end of the other new strand. Called the lagging strand, it is al-

ways growing away from the replication fork. Only short pieces can be synthesized, because if the DNA polymerase were to add continuously to the 3¿ end of that strand, it would need to move far from the replication fork. These 100- to 2000-nucleotide pieces are called Okazaki fragments after their discoverer, Japanese molecular biologist Reiji Okazaki. DNA primase periodically catalyzes the synthesis of an RNA primer on the lagging strand. A separate RNA primer initiates each Okazaki fragment, which DNA polymerase then extends toward the 5¿ end of the previously synthesized fragment. When the RNA primer of the previously synthesized fragment is reached, DNA polymerase degrades and replaces the primer with DNA. The fragments are then joined by DNA ligase, an enzyme that links the 3¿ hydroxyl of one Okazaki fragment to the 5¿ phosphate of the DNA immediately next to it, forming a phosphodiester linkage. (As you will see later in the chapter, DNA ligase also rejoins broken phosphodiester bonds during DNA repair.) DNA synthesis is bidirectional When double-stranded DNA separates, two replication forks form, and the molecule replicates in both directions from the origin of replication. In bacteria, each circular DNA molecule usually has only one origin of replication, so the two replication forks proceed around the circle and eventually meet at the other side to complete the formation of two new DNA molecules. FIGURE 12-16a shows a replicating bacterial plasmid. Plasmids are small, circular DNA

3′ 5′

Leading strand DNA helix

RNA primer

3′ 5′

DNA polymerase

Leading strand is synthesized continuously in direction toward replication fork; lagging strand is synthesized in direction away from replication fork. Both strands require RNA primer for initiation of synthesis because DNA can be elongated only by addition to 3′ end of existing polynucleotide strand.

3′

2

Lagging strand is synthesized as short Okazaki fragments. Okazaki fragment synthesis begins with the synthesis of RNA primer. Note that first Okazaki fragment synthesized is now at far left.

3′

3

After each Okazaki fragment has been elongated by DNA polymerase, RNA primer is degraded, gaps are filled in with DNA, and adjoining fragments are linked by DNA ligase.

5′ Replication fork

3′ 5′

1

3′

Lagging strand (first Okazaki fragment)

3′ 5′

Direction of replication

Leading strand

3′

RNA primers 5′

5′

5′

3′ Two Okazaki fragments

3′ 5′

3′ 5′ Leading strand

3′

DNA ligase

5′ 3′

5′

5′ Third Okazaki fragment

Lagging strand 3′ 5′

FIGURE 12-15 Leading and lagging DNA strands

molecules that carry genes separate from those on a bacterial chromosome. Because plasmids are so small compared to the circular bacterial chromosome, they are possible to clearly photograph during replication. A eukaryotic chromosome consists of one long, linear DNA molecule, so having multiple origins of replication expedites the replication process (FIG. 12-16b). Synthesis continues at each replication fork until it meets a newly synthesized strand coming from the opposite direction. This results in a chromosome containing two DNA double helices, each corresponding to a chromatid.

Enzymes proofread and repair errors in DNA DNA replication occurs only once during each cell generation, and it is important that the process be as accurate as possible to

avoid harmful, or possibly even lethal, mutations. Although base pairing during DNA replication is very accurate, errors do occur. Mechanisms have evolved that ensure that errors in replication are corrected. During replication, DNA polymerases proofread each newly added nucleotide against its template nucleotide. When an error in base pairing is found, DNA polymerase immediately removes the incorrect nucleotide and inserts the correct one. A few uncorrected mutations still occur, but they are very infrequent— on the order of one error for every 109 or 1010 base pairs. When errors have been left uncorrected by the normal repair activities of DNA polymerase during DNA replication, cells make use of other repair mechanisms (although exactly how DNA repair enzymes identify these rare, often-subtle errors in the vast amount of normal DNA is not well understood). In mismatch repair, special enzymes recognize the incorrectly paired nucleotides and

Template DNA (light blue)

New DNA (dark blue) 3′ 5′

5′

Visuals Unlimited, Inc.

3′

50 nm

H. J. Kriegstein and D. S. Hogness, Proc. Nat. Acad. Sci. USA 71 : 135–139, 1974

(a) TEM (above left) of a replicating plasmid of E. coli. Most bacterial plasmids and chromosomes have only one origin of replication. Because DNA synthesis proceeds from that point in both directions, two replication forks form (black arrows), travel around the circle, and eventually meet.

340 nm

5′

Replication “bubbles”

3′

Single replication bubble formed from two merged bubbles

3′ 5′

Replication fork

(b) TEM (left) of two replication forks (black arrows) in a segment of a eukaryotic chromosome that has partly replicated. Eukaryotic chromosome DNA contains multiple origins of replication. DNA synthesis proceeds in both directions from each origin until adjacent replication “bubbles” eventually merge.

FIGURE 12-16 Bidirectional DNA replication in bacteria and eukaryotes The illustrations do not show leading strands and lagging strands.

remove them; DNA polymerases then fill in the missing nucleotides. The observation that individuals with a hereditary defect in a mismatch repair enzyme are likely to develop a type of colon cancer demonstrates the crucial role of mismatch repair enzymes in ensuring the fidelity of DNA replication from one generation to the next. In Chapter 13 you will discover that some types of radiation and chemicals, both in the environment and within the cells themselves, cause mutations in DNA. These mutations are almost always harmful, and cells usually correct mutations by using one or more DNA repair enzymes. About 100 kinds of repair enzymes in the bacterium E. coli and 130 kinds in human cells have been discovered so far. One type of DNA repair—nucleotide excision repair— is commonly used to repair DNA lesions (deformed DNA) caused by the sun’s ultraviolet radiation or by harmful chemicals (FIG. 12-17). Three enzymes are involved in nucleotide excision repair—a nuclease to cut out the damaged DNA, a DNA polymerase

to add the correct nucleotides, and DNA ligase to close the breaks in the sugar–phosphate backbone. Individuals suffering from the disease xeroderma pigmentosum have an inherited defect in a nucleotide excision repair enzyme. Affected individuals develop many skin cancers at an early age because DNA lesions caused by ultraviolet radiation are not repaired.

Telomeres cap eukaryotic chromosome ends Unlike bacterial DNA, which is circular, eukaryotic chromosomes have free ends. Because DNA replication is discontinuous in the lagging strand, DNA polymerases do not complete replication of the strand neatly. At the end of the DNA, a small portion is left unreplicated, and a small, single-strand segment of the DNA is lost with each cell cycle (FIG. 12-18a). The important genetic information is retained because chromosomes have protective end caps, or telomeres, that do not contain protein-coding genes. Telomeres consist of short,

Nuclease enzyme bound to DNA

cell death. Scientists have analyzed cell aging since the 1960s, following the pioneering studies of American biologist Leonard Hayflick, who showed that when normal 5′ 3′ somatic cells of the human body are grown 3′ 5′ in culture, they lose their ability to divide after a limited number of cell divisions. Furthermore, the number of cell divisions 1 A nuclease cuts out a piece of DNA, is determined by the age of the individual including the damaged part. from whom the cells were taken. Cells from a 70-year-old can divide only 20 to 30 times, compared with those from an infant, which can divide 80 to 90 times. Many investigators have observed correlations between telomerase activity and the ability of cells to undergo unlimited 5′ 3′ divisions without showing signs of the ag3′ 5′ ing process. However, they could not find evidence of a causal relationship until the DNA polymerase early 2000s, when scientists at the Geron 2 DNA polymerase and DNA ligase DNA ligase Corporation and the University of Texas complete the repair by synthesizing the Southwestern Medical Center conducted a removed part and sealing the new DNA to the original DNA. direct test. Using recombinant DNA technology (see Chapter 15), they infected New DNA cultured normal human cells with a virus 5′ 3′ that carried the genetic information cod3′ 5′ ing for the catalytic subunit of telomerase. The infected cells not only produced active FIGURE 12-17 Nucleotide excision repair of damaged DNA telomerase, which elongated the telomeres significantly, but also continued to divide noncoding, guanine-rich DNA sequences that repeat many times long past the point at which cell divisions would normally cease. (FIG. 12-18b). For example, in human sperm and egg cells, the seTelomeres and telomerase are an active focus of current requence 5¿¬TTAGGG¬3¿ is repeated more than 1000 times at search, for both scientific and practical reasons. The ability to give the ends of each chromosome. Therefore, although a small amount somatic cells telomerase so they can divide many times more than of telomeric DNA fails to replicate each time the cell divides, a they ordinarily would has many potential therapeutic applications, cell can divide many times before it starts losing essential genetic especially if lost or injured cells must be replaced. However, giving information. such cells the property of unlimited cell division also has potenTelomerase, a special DNA replication enzyme, can lengthen tially serious consequences. For example, if transplanted into the telomeric DNA by adding repetitive nucleotide sequences to the body, cells with active telomerase might behave like cancer cells. ends of eukaryotic chromosomes. This enzyme, which researchCancer cells have the ability to divide hundreds of times in ers discovered in 1984, is typically present in cells that divide an culture; in fact, they are virtually immortal. Most cancer cells, inunlimited number of times, including protozoa and other unicelcluding human cancers of the breast, lung, colon, prostate gland, lular eukaryotes, and most types of cancer cells. In humans and and pancreas, have telomerase, which allows them to maintain telother mammals, active telomerase is usually present in germ line omere length and possibly to resist apoptosis. Research is undercells (which give rise to eggs and sperm) and rapidly dividing cells way to develop anticancer drugs that inhibit telomerase activity. (such as blood cells, cells lining the intestine, and skin cells) but An alternative treatment for cancer is also under study—directing not in most somatic cells of adult tissues. the body’s immune system to recognize and attack cancer cells that When most cells divide for repair or replacement, their chrocontain telomerase. mosome ends shorten. Researchers have demonstrated that in cancer cells of the prostate and pancreas, telomeres are abnormally Review short: they shorten to a critical point, at which time telomerase is reactivated, which may explain the ability of cancer cells to prolif■ How did Meselson and Stahl verify that DNA replication is erate in a rapid and uncontrolled manner. Telomere shortening has semiconservative? also been demonstrated in precancerous lesions from the bladder, ■ Why is DNA replication continuous for one strand but discontinuous breast, colon, esophagus, mouth, and cervix. for the other? Research evidence suggests that the shortening of telomeres ■ How does mismatch repair of DNA compare with nucleotide excision repair? may contribute to cell aging and apoptosis, which is programmed DNA lesion

Unwound DNA before replication 5′

3′

3′

5′ 1

When DNA replication occurs, the 5′ end of each newly synthesized strand contains a short segment of RNA. This RNA functions as a primer on the lagging strand.

5′

3′

3′

5′

+

5′

3′

3′

5′ 2

Removal of the 5′ RNA primer results in DNA molecules with shorter 5′ ends than in the original chromosome. This occurs because there is no way to prime the last section at the 5′ end.

5′

Peter Lansdorp/Visuals Unlimited, Inc.

RNA primer

RNA primer

3′ 5′

3′

+ 5′ 3′

10 μm 3′ 5′

(a) Why one strand of DNA is shorter than that of the preceding generation.

■ ■

(b) LM of duplicated human chromosomes with fluorescent-tagged telomeres (yellow).

FIGURE 12-18 Telomeres

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

12.1 (page 264)

■

1 Summarize the evidence that accumulated during the 1940s and early 1950s demonstrating that DNA is the genetic material. Many early geneticists thought genes were made of proteins. They knew proteins were complex and variable, whereas they thought nucleic acids were simple molecules with a limited ability to store information. ■ Several lines of evidence supported the idea that DNA (deoxyribonucleic acid) is the genetic material. In transformation experiments, the DNA of one strain of bacteria can endow related bacteria with new genetic characteristics. ■ When a bacterial cell becomes infected with a bacteriophage (virus), only the DNA from the virus enters the cell; this DNA is sufficient for the virus to reproduce and form new viral particles. 2 State the questions that these classic experiments addressed: Griffith’s transformation experiment, Avery’s contribution to Griffith’s work, and the Hershey–Chase experiments. ■ Griffith’s transformation experiment addressed this question: Can a genetic trait be transmitted from one bacterial strain to another? (The answer is yes.) Avery’s experiments addressed this question: What molecule is ■ responsible for bacterial transformation? (The answer is DNA.) ■

The Hershey–Chase experiments addressed this question: Is DNA or protein the genetic material in bacterial viruses (phages)? (The answer is DNA.)

12.2 (page 265) 3 Explain how nucleotide subunits link to form a single DNA strand. ■

■

■

Watson and Crick’s model of the structure of DNA demonstrated how information can be stored in the molecule’s structure and how DNA molecules can serve as templates for their own replication. DNA is a polymer of nucleotides. Each nucleotide subunit contains a nitrogenous base, which may be one of the purines (adenine or guanine) or one of the pyrimidines (thymine or cytosine). Each base covalently links to a pentose sugar, deoxyribose, which covalently bonds to a phosphate group. The backbone of each single DNA chain is formed by alternating sugar and phosphate groups, joined by covalent phosphodiester linkages. Each phosphate group attaches to the 5¿ carbon of one deoxyribose and to the 3¿ carbon of the neighboring deoxyribose.

4 Describe how the two strands of DNA are oriented with respect to each other. ■ Each DNA molecule consists of two polynucleotide chains that associate as a double helix. The two chains are antiparallel (running

in opposite directions); at each end of the DNA molecule, one chain has a phosphate attached to a 5¿ deoxyribose carbon, the 5¿ end, and the other has a hydroxyl group attached to a 3¿ deoxyribose carbon, the 3¿ end. Interact with the structure of the DNA double helix by clicking on the figures in CengageNOW.

5 State the base-pairing rules for DNA and describe how complementary bases bind to each other. Hydrogen bonding between specific base pairs holds together the two chains of the helix. Adenine (A) forms two hydrogen bonds with thymine (T); guanine (G) forms three hydrogen bonds with cytosine (C).

■

■

■

A

T

T

A

G

C

C

G

Complementary base pairing between A and T and between G and C is the basis of Chargaff’s rules, which state that A equals T and that G equals C. Because complementary base pairing holds together the two strands of DNA, it is possible to predict the base sequence of one strand if you know the base sequence of the other strand.

12.3 (page 271) 6 Cite evidence from Meselson and Stahl’s experiment that enabled scientists to differentiate between semiconservative replication of DNA and alternative models. ■ When E. coli cells are grown for many generations in a medium containing heavy nitrogen (15N), they incorporate the 15N into their DNA. When researchers transfer cells from a 15N medium to a 14N medium and isolate them after either one or two generations, the density of the DNA in each group is what would be expected if DNA replication were semiconservative. In semiconservative replication, each daughter double helix consists of an original strand from the parent molecule and a newly synthesized complementary strand. 7 Summarize how DNA replicates and identify some unique features of the process. ■ During DNA replication, the two strands of the double helix unwind. Each strand serves as a template for forming a new, complementary strand. Replication is initiated as DNA primase

synthesizes a short RNA primer. DNA polymerase then adds new nucleotide subunits to the growing DNA strand. ■ Additional enzymes and other proteins are required to unwind and stabilize the separated DNA helix. DNA helicases open the double helix, and topoisomerases prevent tangling and knotting. 8 Explain the complexities of DNA replication that make it (a) bidirectional and (b) continuous in one strand and discontinuous in the other. DNA replication is bidirectional, starting at the origin of replica■ tion and proceeding in both directions from that point. A eukaryotic chromosome may have multiple origins of replication and may be replicating at many points along its length at any one time. ■ DNA synthesis always proceeds in a 5¿ ¡ 3¿ direction. This requires that one DNA strand, the lagging strand, be synthesized discontinuously, as short Okazaki fragments. DNA primase synthesizes short RNA primers on the lagging strand, and DNA ligase links Okazaki fragments of newly synthesized DNA. The opposite strand, the leading strand, is synthesized continuously. Watch the process of replication by clicking on the figure in CengageNOW.

9 Discuss how enzymes proofread and repair errors in DNA. During replication, DNA polymerases proofread each newly added nucleotide against its template nucleotide. When an error in base pairing is found, DNA polymerase immediately removes the incorrect nucleotide and inserts the correct one. ■ In mismatch repair, enzymes recognize incorrectly paired nucleotides and remove them; DNA polymerases then fill in the missing nucleotides. ■ Nucleotide excision repair is commonly used to repair DNA lesions caused by the sun’s ultraviolet radiation or by harmful chemicals. Three enzymes are involved: a nuclease to cut out the damaged DNA, a DNA polymerase to add the correct nucleotides, and DNA ligase to close the breaks in the sugar–phosphate backbone. 10 Define telomere, and describe the possible connections between telomerase and cell aging and between telomerase and cancer. ■ Eukaryotic chromosome ends, called telomeres, are short, noncoding, repetitive DNA sequences. Telomeres shorten slightly with each cell cycle but can be extended by the enzyme telomerase. ■ The absence of telomerase activity in certain cells may be a cause of cell aging, in which cells lose their ability to divide after a limited number of cell divisions. ■ Most cancer cells, including human cancers of the breast, lung, colon, prostate gland, and pancreas, have telomerase to maintain telomere length and possibly to resist apoptosis. ■

T E S T YO U R U N D E R S TA N D I N G 1. When Griffith injected mice with a combination of live roughstrain and heat-killed smooth-strain pneumococci, he discovered that (a) the mice were unharmed (b) the dead mice contained living rough-strain bacteria (c) the dead mice contained living smooth-strain bacteria (d) DNA had been transferred from the smooth-strain bacteria to the mice (e) DNA had been transferred from the rough-strain bacteria to the smooth-strain bacteria 2. Which of the following inspired Avery and his colleagues to perform the experiments demonstrating that the transforming factor in bacteria is DNA? (a) the fact that A is equal to T, and G is equal to C (b) Watson and Crick’s model of DNA structure (c) Meselson and Stahl’s studies on DNA replication in E. coli (d) Griffith’s experiments on smooth and rough strains of pneu-

mococci (e) Hershey and Chase’s experiments on the reproduction of bacteriophages 3. In the Hershey–Chase experiment with bacteriophages, (a) harmless bacterial cells permanently transformed into virulent cells (b) DNA was shown to be the transforming factor of earlier bacterial transformation experiments (c) the replication of DNA was conclusively shown to be semiconservative (d) viral DNA was shown to enter bacterial cells and cause the production of new viruses within the bacteria (e) viruses injected their proteins, not their DNA, into bacterial cells 4. The two complementary strands of the DNA double helix are held to each other by (a) ionic bonds between deoxyribose molecules (b) ionic bonds between phosphate groups (c) covalent bonds be-

tween nucleotide bases (d) covalent bonds between deoxyribose molecules (e) hydrogen bonds between nucleotide bases

newly synthesized (e) each double helix consists of one old and one newly synthesized strand

5. If a segment of DNA is 5¿¬CATTAC¬3¿, the complementary DNA strand is (a) 3¿¬CATTAC¬5¿ (b) 3¿¬GTAATG¬5¿ (c) 5¿¬CATTAC¬3¿ (d) 5¿¬GTAATG¬3¿ (e) 5¿¬CATTAC¬5¿

9. Topoisomerases (a) synthesize DNA (b) synthesize RNA primers (c) join Okazaki fragments (d) break and rejoin DNA to reduce torsional strain (e) prevent single DNA strands from joining to form a double helix

6. Each DNA strand has a backbone that consists of alternating (a) purines and pyrimidines (b) nucleotide bases (c) hydrogen bonds and phosphodiester linkages (d) deoxyribose and phosphate (e) phosphate and phosphodiester linkages

10. A lagging strand forms by (a) joining primers (b) joining Okazaki fragments (c) joining leading strands (d) breaking up a leading strand (e) joining primers, Okazaki fragments, and leading strands 11. The immediate source of energy for DNA replication is (a) the hydrolysis of nucleotides with three phosphate groups (b) the oxidation of NADPH (c) the hydrolysis of ATP (d) electron transport (e) the breaking of hydrogen bonds

7. The experiments in which Meselson and Stahl grew bacteria in heavy nitrogen conclusively demonstrated that DNA (a) is a double helix (b) replicates semiconservatively (c) consists of repeating nucleotide subunits (d) has complementary base pairing (e) is always synthesized in the 5¿ ¡ 3¿ direction 8. The statement “DNA replicates by a semiconservative mechanism” means that (a) only one DNA strand is copied (b) first one DNA strand is copied, and then the other strand is copied (c) the two strands of a double helix have identical base sequences (d) some portions of a single DNA strand are old, and other portions are

12. Which of the following statements about eukaryotic chromosomes is false? (a) eukaryotic chromosomes have free ends (b) telomeres contain protein-coding genes (c) telomerase lengthens telomeric DNA (d) telomere shortening may contribute to cell aging (e) cells with active telomerase may undergo many cell divisions

CRITICAL THINKING 1. What characteristics must a molecule have to function as genetic material?

6. EVOLUTION LINK. How does the fact that DNA is the universal molecule of inheritance in all cells support evolutionary theory?

2. What important features of the structure of DNA are consistent with its role as the chemical basis of heredity? 3. What do telomeres on the ends of chromosomes protect against? 4. ANALYZING DATA. In the Hershey–Chase experiment, the radioactive label 32P was present inside bacterial cells (i.e., in the pellet), whereas the radioactive label 35S was present outside bacterial cells (in the supernatant). What would the researchers have concluded had the reverse been true—that is, if the radioactive label 35S was inside the cells and the radioactive label 32P was outside the cells? 5. ANALYZING DATA. In the following diagram of DNA replication, label the 3¿ and 5¿ ends of the leading strand, lagging strand, and the two strands of the DNA double helix. Leading strand DNA helix RNA primer DNA polymerase Replication fork Lagging strand (first Okazaki fragment)

Direction of replication

Additional questions are available in CengageNOW at www.cengage.com/ login.

13

Gene Expression

Professor Oscar Miller/Science Photo Library/Photo Researchers, Inc.

Visualizing transcription. In this TEM, RNA molecules (lateral strands) are synthesized as complementary copies of a DNA template (central axis).

KEY CONCEPTS

I

n Chapter 12, we discussed how the cell accurately replicates the nucleotide sequence in DNA to pass the genetic material unaltered

13.1 Beadle and Tatum demonstrated the relationship

to the next generation. Watson and Crick originally described the basic

between genes and proteins in the 1940s.

features of the DNA double helix, which scientists now know to be the

13.2 The transmission of information in cells is typically from DNA to RNA to polypeptide.

same in all cells studied to date—from the simplest bacterial cells to

13.3 A sequence of DNA base triplets is transcribed into mRNA codons.

13.4 A sequence of mRNA codons is translated into a sequence of amino acids in a polypeptide.

13.5 Bacterial and eukaryotic cells differ in certain details of gene expression.

13.6 Mutations can cause changes in phenotype.

intricate human cells. By the mid-1950s, researchers had determined that the sequence of bases in DNA contains the information that specifies all the proteins the cell needs. However, more than a decade of intense investigation by many scientists preceded a fundamental understanding of how cells convert DNA information into the amino acid sequences of proteins. Much of that understanding came from studying the functions of bacterial genes. After Watson and Crick discovered the structure of DNA, researchers

chose bacterial cells as the organisms for these studies because

Tyrosine

bacteria grow quickly and easily and because they contain the minimal amount of DNA needed for growth and reproduction. As researchers learned that all organisms have fundamental genetic similarities, the validity, as well as the utility, of this approach was repeatedly confirmed.

Functional enzyme absent

Homogentisic acid

Functional enzyme present

In this chapter we examine the evidence that accumulated in the first half of the 20th century indicating that most genes specify

Disease condition

Normal metabolism

the structure of proteins. We then consider how DNA affects the

ALKAPTONURIA

Maleylacetoacetate

phenotype of the organism at the molecular level through the process of gene expression. Gene expression involves a series of steps in which the information in the sequence of bases in DNA specifies the makeup of the cell’s proteins. The proteins affect the pheno-

Homogentisic acid excreted in urine; turns black when exposed to air

type in some way; these effects range from readily observable

CO2 H2O

physical traits to subtle changes detectable only at the biochemical level. The first major step of gene expression is transcription, the synthesis of RNA molecules complementary to the DNA (see photograph). The second major step is translation, in which RNA becomes a coded template to direct polypeptide synthesis.

13.1 DISCOVERY OF THE GENE– PROTEIN RELATIONSHIP ■ ■ LEARNING OBJECTIVES 1 2

Summarize the early evidence indicating that most genes specify the structure of proteins. Describe Beadle and Tatum’s experiments with Neurospora.

The idea that genes and proteins are connected originated early in the 20th century, shortly after scientists rediscovered Mendel’s principles. In the first edition of his book, Inborn Errors of Metabolism (1908), Archibald Garrod, a physician and biochemist, discussed a rare genetic disease called alkaptonuria, which scientists hypothesized had a simple recessive inheritance pattern. The condition involves the metabolic pathway that breaks down the amino acid tyrosine, ultimately converting it to carbon dioxide and water. An intermediate in this pathway, homogentisic acid, accumulates in the urine of affected people, turning it black when exposed to air (FIG. 13-1). Other symptoms of alkaptonuria include the later development of arthritis and, in men, stones in the prostate gland. In Garrod’s time, scientists knew about enzymes but did not recognize they were proteins. Garrod hypothesized that people with alkaptonuria lack the enzyme that normally oxidizes homogentisic acid. Before the second edition of his book had been published, in 1923, researchers found that affected people do indeed lack the enzyme that oxidizes homogentisic acid. Garrod’s hypothesis was correct: a mutation in this gene is associated with the absence of a specific enzyme. Shortly thereafter, in 1926, U.S. biochemist James Sumner purified a different enzyme, urease, and showed it to be a protein. This was the first clear identification of an enzyme as a protein. In 1946, Sumner was awarded the Nobel Prize in Chemistry for being the first to crystallize an enzyme.

FIGURE 13-1 An “inborn error of metabolism” Garrod proposed that alkaptonuria is a genetic disease caused by the absence of the enzyme homogentisic acid oxidase, which normally converts homogentisic acid to maleylacetoacetate. Thus, the acid accumulates in the blood and is excreted in the urine, which turns black on contact with the air.

Beadle and Tatum proposed the one-gene, one-enzyme hypothesis A major advance in understanding the relationship between genes and enzymes came in the early 1940s, when U.S. geneticists George Beadle and Edward Tatum and their associates developed a new approach to the problem. Most efforts until that time had focused on studying known phenotypes, such as eye color in Drosophila or pigments in plants, and seeking to determine what biochemical reactions they affected. Researchers found that a series of biosynthetic reactions controls specific phenotypes, but it was not clear whether the genes themselves were acting as enzymes or whether they controlled the workings of enzymes in some indirect way. Beadle and Tatum decided to take the opposite approach. Rather than try to identify the enzymes affected by single genes, they decided to look for mutations interfering with known metabolic reactions that produce essential molecules, such as amino acids and vitamins. They chose as an experimental organism a fungus, the pink-orange bread mold Neurospora, for several important reasons. Wild-type Neurospora is easy to grow in culture. (The adjective wild-type refers to an individual with the normal phenotype.) Neurospora manufactures all its essential biological molecules when grown on a simple growth medium (minimal medium) containing only sugar, salts, and the vitamin biotin. However, a mutant Neurospora strain that cannot make a substance such as an amino acid can still grow if that substance is added to the growth medium. Neurospora is also an ideal experimental organism because it grows primarily as a haploid organism. The haploid condition allows a researcher to immediately identify a recessive mutant allele; there is no homologous chromosome that might carry a dominant allele that would mask its expression.

Beadle and Tatum began by exposing thousands of haploid wild-type Neurospora asexual spores to X-rays or ultraviolet radiation to produce mutant strains. They first grew each irradiated strain on a complete growth medium, which contained all the amino acids and vitamins normally made by Neurospora. Then they tested each strain on the minimal medium described previously. About 1% to 2% of the strains that grew on the complete medium failed to grow after transfer to the minimal medium. Beadle and Tatum reasoned that such strains carried a mutation that prevented the fungus from producing a chemical essential for growth. Further testing of the mutant strain on media containing different combinations of amino acids, vitamins, and other nutrients enabled the investigators to determine the exact compound required (FIG. 13-2). The work on Neurospora revealed that each mutant strain had a mutation in only one gene and that each gene affected only one enzyme. Beadle and Tatum stated this one-to-one correspondence between genes and enzymes as the one-gene, one-enzyme hypothesis. In 1958, they received the Nobel Prize in Physiology or Medicine for discovering that genes regulate specific chemical events. The work of Beadle, Tatum, and others led to a more precise understanding of what a gene is and to additional predictions about the chemical nature of genes. The idea that a gene encodes the information to produce a single enzyme held for almost a decade, until additional findings required a modification of this definition. In the late 1940s, researchers began to understand that genes control not only enzymes but other proteins as well. The U.S. chemist Linus Pauling and his colleagues demonstrated in 1949 that a mutation of a single gene alters the structure of hemoglobin. This particular mutant form of hemoglobin is associated with the genetic disease sickle cell anemia (discussed in Chapter 16). British biochemist Vernon Ingram extended Pauling’s research in 1957 when he determined that sickle cell hemoglobin and normal hemoglobin differ by only one amino acid. Studies by other scientists showed that many proteins are constructed from two or more polypeptide chains, each of which is under the control of a different locus. For example, hemoglobin contains two types of polypeptide chains, the a and b subunits. See the structure of hemoglobin in Figure 3-22a. Sickle cell anemia results from a mutation affecting the b subunits. Scientists therefore extended the definition of a gene to include that one gene is responsible for one polypeptide chain. Even this definition has proved only partially correct, although as you will see later in this chapter, scientists still define a gene in terms of its product. Although the elegant work of Beadle and Tatum and others demonstrated that genes are expressed in the form of proteins, the mechanism of gene expression was completely unknown. After Watson and Crick’s discovery of the structure of DNA, many scientists worked to understand exactly how gene expression takes place. We begin with an overview of gene expression and then consider the various steps in more detail.

Review ■ ■

What is the one-gene, one-enzyme hypothesis? What were the contributions of each of the following scientists— Garrod, Beadle and Tatum, and Pauling—to our understanding of the relationship between genes and proteins?

KEY EXPERIMENT QUESTION: Can genes code for enzymes? HYPOTHESIS: Mutations induced in Neurospora, a type of bread mold, correspond to the absence of functional enzymes. EXPERIMENT: Beadle and Tatum irradiated Neurospora spores to induce mutations. They grew cultures derived from these spores on complete growth medium containing all amino acids, vitamins, and other nutrients that Neurospora normally makes for itself. They identified nutritional requirements in mutant strains by testing for growth on minimal media supplemented with individual vitamins or amino acids. Expose Neurospora spores to UV light or X-rays.

Fungal growth (mycelium)

Each irradiated spore is used to establish culture on complete growth medium (minimal medium plus amino acids, vitamins, etc.).

RESULTS AND CONCLUSION: They transferred some fungal cells to each of three tubes: one with minimal medium plus vitamins; one with minimal medium plus amino acids; and one with minimal medium (the control). In this example the mutant fungus grew on the minimal medium only if it was supplemented with amino acids.

Transfer cells to minimal medium plus vitamins.

Transfer cells to minimal medium plus amino acids.

Transfer cells to minimal medium (control).

To determine which amino acid the mutant fungus had lost the ability to synthesize, they transferred the mutant fungus to tubes containing minimal medium and one amino acid only.

Minimal medium plus arginine

Minimal medium plus tryptophan

Minimal medium plus lysine

Minimal medium plus leucine

Minimal medium plus other amino acids

In this example, only the medium containing the amino acid arginine supported growth, indicating that the mutation affected some part of arginine’s biosynthetic pathway. Beadle and Tatum then identified the enzyme-catalyzed step that was blocked in the synthesis of arginine. On the basis of many similar experiments, they concluded that each gene controls the production of a single enzyme.

FIGURE 13-2 The Beadle–Tatum experiments

13.2 INFORMATION FLOW FROM DNA TO PROTEIN: AN OVERVIEW

5′

O

–O

P

4 5

Outline the flow of genetic information in cells, from DNA to RNA to polypeptide. Compare the structures of DNA and RNA. Explain why the genetic code is said to be redundant and virtually universal, and discuss how these features may reflect its evolutionary history.

Although the sequence of bases in DNA determines the sequence of amino acids in polypeptides, cells do not use the information in DNA directly. Instead, a related nucleic acid, ribonucleic acid (RNA), links DNA and protein. When a gene that codes for a protein is expressed, first an RNA copy is made of the information in the DNA. It is this RNA copy that provides the information that directs polypeptide synthesis. Like DNA, RNA is a polymer of nucleotides, but it has some important differences (FIG. 13-3). RNA is usually single-stranded, although internal regions of some RNA molecules may have complementary sequences that allow the strand to fold back and pair to form short, double-stranded segments. As shown in Figure 13-3, the sugar in RNA is ribose, which is similar to deoxyribose of DNA but has a hydroxyl group at the 2¿ position. (Compare ribose with the deoxyribose of DNA, shown in Figure 12-4, which has a hydrogen at the 2¿ position.) The base uracil substitutes for thymine and, like thymine, is a pyrimidine that can form two hydrogen bonds with adenine. Hence, uracil and adenine are a complementary pair.

DNA is transcribed to form RNA The process by which RNA is synthesized resembles DNA replication in that the sequence of bases in the RNA strand is determined by complementary base pairing with one of the DNA strands, the template strand (FIG. 13-4). Because RNA synthesis takes the information in one kind of nucleic acid (DNA) and copies it as another nucleic acid (RNA), this process is called transcription (“copying”). Three main kinds of RNA molecules are transcribed: messenger RNA, transfer RNA, and ribosomal RNA. Messenger RNA (mRNA) is a single strand of RNA that carries the information for making a protein. Each of the 45 or so kinds of transfer RNAs (tRNAs) is a single strand of RNA that folds back on itself to form a specific shape. Each kind of tRNA bonds with only one kind of amino acid and carries it to the ribosome. (Because there are more kinds of tRNA molecules than there are amino acids, many amino acids are carried by two or more kinds of tRNA molecules.) Ribosomal RNA (rRNA), which is in a globular form, is an important part of the structure of ribosomes and has catalytic functions needed during protein synthesis.

RNA is translated to form a polypeptide Following transcription, the transcribed information in the mRNA is used to specify the amino acid sequence of a polypeptide (see

O

O

■ ■ LEARNING OBJECTIVES 3

O

CH2

N

O

N

Ribose O –O

Uracil

O

OH O

P

NH2 N

O CH2

N

N

O

N Adenine

Ribose O –O

OH

NH2

O

P

N

O CH2 O

O Cytosine

N

Ribose O –O

O

OH O

N

O CH2 O

N

P

Ribose OH 3′

N N Guanine

NH2

OH

FIGURE 13-3 The nucleotide structure of RNA The nucleotide subunits of RNA are joined by 5¿ ¡ 3¿ phosphodiester linkages, like those found in DNA. Adenine, guanine, and cytosine are present, as in DNA, but the base uracil replaces thymine. All four nucleotide types contain the five-carbon sugar ribose, which has a hydroxyl group (blue) on its 2¿ carbon atom.

Fig. 13-4). This process is called translation because it involves conversion of the “nucleic acid language” in the mRNA molecule into the “amino acid language” of protein. In translation of the genetic instructions to form a polypeptide, a sequence of three consecutive bases in mRNA, called a codon, specifies one amino acid. For example, one codon that corresponds to the amino acid phenylalanine is 5¿¬UUC¬3¿. Because each codon consists of three nucleotides, the code is described as a triplet code. The assignments of codons for amino acids and for start and stop signals are collectively named the genetic code (FIG. 13-5).

KEY POINT

transcription translation Protein synthesis requires two major steps: DNA ¬¬ ¡ RNA ¬¬¡ protein

Nontemplate strand A G 3′

5′ DNA

A

3′

T

mRNA (complementary copy of template DNA strand)

Polypeptide

C G A A G C T T T

5′

Transcription

G U U C C A A G

3′

C U G A A G C T T C

G A

Template strand

A U G A C U U G C G A A U G U U U C 5′

H2N

Codon 1

Codon 2

Codon 3

Codon 4

Codon 5

Codon 6

Met

Thr

Cys

Glu

Cys

Phe

O C

OH

Translation

FIGURE 13-4 An overview of transcription and translation In transcription, messenger RNA is synthesized as a complementary copy of one of the DNA strands, the template strand. Messenger RNA carries genetic information in the form of sets of three bases, or codons, each of which specifies one amino acid. Codons are translated consecutively,

U

UUU Phe UUC UUA Leu UUG

UCU UCC Ser UCA UCG

UAU Tyr UAC UAA Stop UAG Stop

UGU Cys UGC UGA Stop UGG Trp

U C A G

C

CUU CUC Leu CUA CUG

CCU CCC Pro CCA CCG

CAU His CAC CAA Gln CAG

CGU CGC Arg CGA CGG

U C A G

A

AUU AUC Ile AUA AUG Met or start

ACU ACC Thr ACA ACG

AAU Asn AAC AAA Lys AAG

AGU Ser AGC AGA Arg AGG

U C A G

G

GUU GUC Val GUA GUG

GCU GCC Ala GCA GCG

GAU Asp GAC GAA Glu GAG

GGU GGC Gly GGA GGG

U C A G

= Stop codon = Start codon

Transfer RNAs are crucial parts of the decoding machinery because they act as “adapters” that connect amino acids and nucleic acids. This mechanism is possible because each tRNA can (1) link with a specific amino acid and (2) recognize the appropriate mRNA codon for that particular amino acid (FIG. 13-6). A particular tRNA can recognize a particular codon because it has a sequence of three bases, called the anticodon, that hydrogen-bonds with the mRNA codon by complementary base pairing. The exact anticodon that is complementary to the codon for phenylalanine in our example is 3¿¬AAG¬5¿. Translation requires that each tRNA anticodon be hydrogenbonded to the complementary mRNA codon and that the amino acids carried by the tRNAs be linked in the order specified by the sequence of codons in the mRNA. Ribosomes, the site of translation, are organelles composed of two different subunits, each containing protein and rRNA. (The structure and function of ribosomes were introduced in Chapter 4.) Ribosomes attach to the 5¿ ▲

First letter (5′ end)

G

Third letter (3′ end)

Second letter C A

U

thus specifying the linear sequence of amino acids in the polypeptide chain. Translation requires tRNA and ribosomes (not shown). The figure depicts transcription and translation in bacteria. In eukaryotes, transcription takes place in the nucleus and translation occurs in the cytosol.

FIGURE 13-5 The genetic code

The genetic code specifies all possible combinations of the three bases that compose codons in mRNA. Of the 64 possible codons, 61 specify amino acids (see Fig. 3-17 for an explanation of abbreviations). The codon AUG specifies the amino acid methionine and also signals the ribosome to initiate translation (“start”). Three codons—UAA, UGA, and UAG—do not specify amino acids; they terminate polypeptide synthesis (“stop”).

OH 3′ end

Amino acid accepting end 3′

Loop 3

5′

Hydrogen bonds Loop 3 U C Loop 1

G C

T

C A C A G

P 5′ end G C G G A U U U

H H

C U C A

A

Modified nucleotides

C G A G A

C G A G G U C

Loop 2

C C A G A

O C

G

O 3′

5′

G G

G

A A G

A

C

A

G

U

Loop 2 A

N

Amino acid (phenylalanine)

Loop 1

G UG U U

Anticodon

A C C A C G C U U A A

Anticodon

Anticodon

(a) The 3-D shape of a tRNA molecule is determined by hydrogen bonds formed between complementary bases.

(b) One loop contains the anticodon; these unpaired bases pair with a complementary mRNA codon. The amino acid attaches to the terminal nucleotide at the hydroxyl (OH) 3′ end.

FIGURE 13-6 Three representations of a tRNA molecule

end of the mRNA and travel along it, allowing the tRNAs to attach sequentially to the codons of mRNA. In this way the amino acids carried by the tRNAs take up the proper position to be joined by peptide bonds in the correct sequence to form a polypeptide.

Biologists cracked the genetic code in the 1960s Before the genetic code was deciphered, scientists had become interested in how a genetic code might work. The Watson and Crick model of DNA showed it to be a linear sequence of four different nucleotides. If each nucleotide coded for a single amino acid, the genetic code could specify only 4 amino acids, not the 20 found in the vast variety of proteins in the cell. Scientists saw that the DNA bases could serve as a four-letter “alphabet” and hypothesized that three-letter combinations of the four bases would make it possible to form a total of 64 “words,” more than enough to specify all the naturally occurring amino acids. In 1961, Crick and British scientist Sydney Brenner concluded from experimental evidence that the code used nonoverlapping triplets of bases. They predicted that the code is read, one triplet at a time, from a fixed starting point that establishes the reading frame for the genetic message. Marshall Nirenberg, a U.S. biochemist, and his postdoctoral researcher, Heinrich Matthaei, studied protein synthesis outside living cells in purified cell-free systems derived from the bacterium

(c) This stylized diagram of an aminoacyl-tRNA shows that the amino acid attaches to tRNA by its carboxyl group, leaving its amino group exposed for peptide bond formation.

Escherichia coli. Nirenberg and Matthaei obtained the first experimental evidence indicating the assignment of triplets to specific amino acids. By constructing artificial mRNA molecules with known base sequences, they determined which amino acids would be incorporated into protein. For example, when they added the synthetic mRNA polyuridylic acid (UUUUUUUUU . . .) to a mixture of purified ribosomes, aminoacyl-tRNAs (amino acids linked to their respective tRNAs), and essential cofactors needed to synthesize polypeptide, only phenylalanine was incorporated into the resulting polypeptide chain. The inference was inescapable that the UUU triplet codes for phenylalanine. Similar experiments showed that polyadenylic acid (AAAAAAAAA . . .) codes for a polypeptide of lysine and that polycytidylic acid (CCCCCCCCC . . .) codes for a polypeptide of proline. By using mixed nucleotide polymers (such as a random polymer of A and C) as artificial messengers, researchers such as H. Gobind Khorana, then at the University of Wisconsin, assigned the other codons to specific amino acids. However, three of the codons— UAA, UGA, and UAG—did not specify any amino acids. These codons are now known to be the signals that terminate the coding sequence for a polypeptide chain. By 1967, the genetic code was completely “cracked,” and scientists had identified the coding assignments of all 64 possible codons shown in Figure 13-5. In 1968, Nirenberg and Khorana received the Nobel Prize in Physiology or Medicine for their work in deciphering the genetic code.

Scientists have discovered some minor exceptions to the universality of the genetic code. In several unicellular protozoa, UAA and UGA code for the amino acid glutamine instead of functioning as stop signals. Other exceptions are found in mitochondria, which contain their own DNA and protein synthesis machinery for some genes. These slight coding differences vary with the organism, but keep in mind that in each case, all the other coding assignments are identical to those of the standard genetic code.

Remember that the genetic code is an mRNA code. The tRNA anticodon sequences, as well as the DNA sequence from which the message is transcribed, are complementary to the sequences in Figure 13-5. For example, the mRNA codon for the amino acid methionine is 5¿¬AUG¬3¿. It is transcribed from the DNA base sequence 3¿¬TAC¬5¿, and the corresponding tRNA anticodon is 3¿¬UAC¬5¿.

The genetic code is virtually universal The genetic code is redundant

Perhaps the single most remarkable feature of the code is that it is essentially universal. Over the years, biologists have examined the genetic code in a diverse array of species and found it the same in organisms as different as bacteria, redwood trees, jellyfish, and humans (FIG. 13-7). These findings strongly suggest that the code is an ancient legacy that evolved early in the history of life (discussed in Chapter 21).

Given 64 possible codons and only 20 common amino acids, it is not surprising that more than one codon specifies certain amino acids. This redundancy in the genetic code has certain characteristic patterns. The codons CCU, CCC, CCA, and CCG are “synonymous” in that they all code for the amino acid proline. The only difference among the 4 codons involves the nucleotide at the 3¿ end of the triplet. Although the code may be read three nucleotides at a time, only the first two nucleotides seem to contain specific information for proline. A similar pattern can be seen for several other amino acids. Only methionine and tryptophan are specified by single codons. Each of the other amino acids is specified by 2 to 6 different codons. Crick first proposed this apparent breach of the base-pairing rules as the wobble hypothesis. He reasoned that the third nucleotide of a tRNA anticodon (which is the 5¿ base of the anticodon sequence) may sometimes form hydrogen bonds with more than one kind of third nucleotide (the 3¿ base) of a codon. Investigators later established this experimentally by determining the anticodon sequences of tRNA molecules and testing their specificities in artificial systems. Some tRNAs bond exclusively to one codon, but other tRNA molecules pair with as many as three separate codons that differ in their third nucleotide but that specify the same amino acid. Thus, the wobble hypothesis accounts for the possible variation in base pairing between the third base of a codon and the corresponding base in its anticodon. “Wobble” results in several acceptable forms of base pairing between mRNA and tRNA.

Review ■

Keith V. Wood/Visuals Unlimited

■

13.3 TRANSCRIPTION ■ ■ LEARNING OBJECTIVES 6 Compare the processes of transcription and DNA replication, identifying 7

FIGURE 13-7 Genetically engineered tobacco plant This plant glows as it expresses the luciferase gene from a firefly. Luciferase is an enzyme that catalyzes a reaction that produces a flash of light. This classic experiment indicated that animal genes can be expressed in plants.

Sketch a simple flow diagram that shows the relationships among the following: RNA, translation, DNA, transcription, and polypeptide. How are the structures of DNA and RNA similar? How are they different?

both similarities and differences. Compare bacterial and eukaryotic mRNAs, and explain the functional significance of their structural differences.

Now that we have presented an overview of information flow from DNA to RNA to polypeptide, let us examine the entire pro-

cess more closely. The first stage in the flow of information from DNA to polypeptide is the transcription of a DNA nucleotide sequence into an RNA nucleotide sequence. Stanford University biologist Roger Kornberg painstakingly worked out the details of transcription, for which he was awarded the 2006 Nobel Prize in Chemistry. In eukaryotic transcription, most RNA synthesis requires one of three RNA polymerases, enzymes present in all cells. The three RNA polymerases differ in the kinds of RNA synthesis they catalyze. RNA polymerase I catalyzes the synthesis of several kinds of rRNA molecules that are components of the ribosome; RNA polymerase II catalyzes the production of the protein-coding mRNA; and RNA polymerase III catalyzes the synthesis of tRNA and one of the rRNA molecules. RNA polymerases require DNA as a template and have many similarities to the DNA polymerases discussed in Chapter 12. Like DNA polymerases, RNA polymerases carry out synthesis in the 5¿ ¡ 3¿ direction; that is, they begin at the 5¿ end of the RNA molecule being synthesized and then continue to add nucleotides at the 3¿ end until the molecule is complete (FIG. 13-8). RNA polymerases use nucleotides with three phosphate groups (for example, ATP and GTP) as substrates for the polymerization reaction. As the nucleotides link to the 3¿ end of the RNA, two of the phosphates are removed. These reactions are strongly exergonic and do not need the input of additional energy. Recall from Chapter 12 that whenever nucleic acid molecules associate by complementary base pairing, the two strands are antiparallel. Just as the two paired strands of DNA are antiparallel, the template strand of the DNA and the complementary RNA strand are also antiparallel. Therefore, when transcription takes place, as RNA is synthesized in its 5¿ ¡ 3¿ direction, the DNA template is read in its 3¿ ¡ 5¿ direction. Scientists conventionally refer to a sequence of bases in a gene or the mRNA sequence transcribed from it as upstream or downstream from some reference point. Upstream is toward the 5¿ end of the mRNA sequence or the 3¿ end of the template DNA strand. Downstream is toward the 3¿ end of the RNA or the 5¿ end of the template DNA strand. Upstream 5′—A—T—G—A—C—T—3′ 3′—T—A—C—T—G—A—5′

Downstream nontemplate DNA strand template DNA strand

Direction of transcription 5′—A—U— G—A— C—U—3′

RNA

The synthesis of mRNA includes initiation, elongation, and termination In both bacteria and eukaryotes, the nucleotide sequence in DNA to which RNA polymerase and associated proteins initially bind is called the promoter. Because the promoter is not transcribed, RNA polymerase moves past the promoter to begin transcription of the protein-coding sequence of DNA. Different genes may have

O

Growing RNA O strand P

Template DNA strand

5′ end O O

O

3′ direction

P O

O –O

P

O

O

O

O CH2

U O

–O

P

H H

A

CH2 O

OH O

O CH2

O G

OH OH

H H H

P

CH2 O O

O–

O

CH2

A

H H

O–

P O

T CH2

OH OH

O O

Nucleotide added to 3′ end growing chain by RNA polymerase

O–

P O

C

O O

O–

P

P

O–

G 5′ direction

FIGURE 13-8 Animated A molecular view of transcription Incoming nucleotides with three phosphates pair with complementary bases on the template DNA strand (right). RNA polymerase cleaves two phosphates (not shown) from each nucleotide and covalently links the remaining phosphate to the 3¿ end of the growing RNA chain. Thus, RNA, like DNA, is synthesized in the 5¿ ¡ 3¿ direction.

slightly different promoter sequences, so the cell can direct which genes are transcribed at any one time. Bacterial promoters are usually about 40 bases long and lie in the DNA just upstream of the point at which transcription will begin. Once RNA polymerase has recognized the correct promoter, it unwinds the DNA double helix and initiates transcription (FIG. 13-9). Unlike DNA synthesis, RNA synthesis does not require a primer. However, transcription requires several proteins in addition to RNA polymerase; we discuss these in Chapter 14. The first nucleotide at the 5¿ end of a new mRNA chain initially retains its triphosphate group (see Fig. 13-8). However, during the elongation stage of transcription, as each additional nucleotide is incorporated at the 3¿ end of the growing RNA molecule, two of its phosphates are removed in an exergonic reaction that leaves the remaining phosphate to become part of the sugar–phosphate backbone (as in DNA replication). The last nucleotide to be incorporated has an exposed 3¿-hydroxyl group. Elongation of RNA continues until termination, when RNA polymerase recognizes a termination sequence consisting of a

KEY POINT

The mRNA is synthesized in the 5¿ to 3¿ direction as the template strand of the DNA molecule is read in the 3¿ to 5¿ direction. RNA polymerase binds to promoter region in DNA

5′ 3′

1

Initiation. RNA polymerase unwinds DNA double helix and initiates RNA synthesis.

3′ 5′

2

Elongation. Additional nucleotides are added to the 3′ end of RNA molecule. DNA double helix re-forms following transcription.

3′ 5′

3

Elongation continues.

3′ DNA 5′

4

Termination. RNA polymerase recognizes termination sequence. RNA transcript and RNA polymerase are released.

Termination sequence

Promoter region Direction of transcription 5′ 3′

3′ 5′ DNA

DNA template strand 3′

5′ RNA transcript

Rewinding of DNA 5′ 3′

5′ 3′

Unwinding of DNA 3′

5′

5′

3′ RNA transcript RNA polymerase

FIGURE 13-9 Animated An overview of transcription: initiation, elongation, and termination

specific sequence of bases in the DNA template. This signal leads to a separation of RNA polymerase from the template DNA and the newly synthesized RNA. Termination of transcription occurs by different mechanisms in bacteria and eukaryotes. In bacteria, transcription stops at the end of the termination sequence. When RNA polymerase comes to the termination sequence, it releases the DNA template and the new RNA strand. In eukaryotic cells, RNA polymerase adds nucleotides to the mRNA molecule after it passes the termination sequence. The 3¿ end of the mRNA becomes separated from RNA polymerase about 10 to 35 nucleotides past the termination sequence. Only one strand in a protein-coding region of DNA is used as a template. For example, consider a segment of DNA that contains the following DNA base sequence in the template strand: 3¿¬TAACGGTCT¬5¿

If the complementary DNA strand 5¿¬ATTGCCAGA¬3¿

were used as a template, a message would be produced specifying an entirely different (and probably nonfunctional) protein. However, just because one strand is transcribed for a given gene does not mean the same DNA strand is always the template for all genes throughout the length of a chromosome-sized DNA molecule. Instead, a particular strand may serve as the template strand for some genes and the nontemplate strand for others (FIG. 13-10).

Messenger RNA contains base sequences that do not directly code for protein A completed mRNA molecule contains more than the nucleotide sequence that codes for a protein. FIGURE 13-11 shows a typical bacterial mRNA. (The unique features of eukaryotic mRNA are discussed in the next section.) In both bacteria and eukaryotes, RNA polymerase starts transcription of a gene upstream of the proteincoding DNA sequence. As a result, the mRNA has a noncoding leader sequence at its 5¿ end. The leader has recognition sites for ribosome binding, which properly position the ribosomes to translate the message. The start codon follows the leader sequence and signals the beginning of the coding sequence that contains the

Promoter region

mRNA transcript 5′ Gene 2

RNA polymerase

5′

mRNA transcript 5′

Promoter region

3′

3′

3′

Promoter region

5′

3′

3′ Gene 1

Gene 3 5′ mRNA transcript

FIGURE 13-10 The template strand for one gene may be the nontemplate strand for another gene Only one of the two strands is transcribed for a given gene, but the opposite strand may be transcribed for a neighboring gene. Each transcript starts at its own promoter region (orange). The orange arrow associated with each promoter region indicates the direction of transcription.

actual message for the polypeptide. Unlike coding in eukaryotic cells, in bacterial cells it is common for more than one polypeptide to be encoded by a single mRNA molecule (see Chapter 14). At the end of each coding sequence, a stop codon signals the end of the protein. The stop codons—UAA, UGA, and UAG—end both bacterial and eukaryotic messages. They are followed by noncoding 3¿ trailing sequences, which vary in length.

Eukaryotic mRNA is modified after transcription and before translation Other differences exist between bacterial and eukaryotic mRNA. Bacterial mRNAs are used immediately after transcription, without further processing. In eukaryotes, the original transcript, called precursor mRNA, or pre-mRNA, is modified in several ways while it is still in the nucleus. These posttranscriptional modification and processing activities produce mature mRNA for transport and translation. Modification of the eukaryotic message begins when the growing RNA transcript is about 20 to 30 nucleotides long. At that point, enzymes add a 5¿ cap to the 5¿ end of the mRNA chain (FIG. 13-12). Promoter region

The cap is in the form of an unusual nucleotide, 7-methylguanosine, which is linked to the mRNA transcript by three phosphate groups. Eukaryotic ribosomes cannot bind to an uncapped message. Capping may also protect the mRNA from certain types of degradation and may therefore partially explain why eukaryotic mRNAs are much more stable than bacterial mRNAs. Eukaryotic mRNAs have half-lives ranging from 30 minutes to as long as 24 hours. The average half-life of an mRNA molecule in a mammalian cell is about 10 hours compared with 2 minutes in a bacterial cell. A second modification of eukaryotic mRNA, known as polyadenylation, may occur at the 3¿ end of the molecule. Near the 3¿ end of a completed message usually lies a sequence of bases that serves as a signal for adding many adenine-containing nucleotides, known as a poly-A tail (for polyadenylated). Enzymes in the nucleus recognize the signal for the poly-A tail and cut the mRNA molecule at that site. This process is followed by the enzymatic addition of a string of 100 to 250 adenine nucleotides to the 3¿ end. Polyadenylation appears to have several functions: it helps export the mRNA from the nucleus, stabilizes some mRNAs against degradation in the cytosol (the longer the poly-A tail, the longer

Transcribed region

mRNA termination sequence

DNA 5′

5′

3′

3′ Upstream leader sequences

Protein-coding sequences Translated region Start codon

Downstream trailing sequences

Stop codon

mRNA O– P –O– P –O– P –O–

–OH

5′ end

3′ end

Polypeptide

FIGURE 13-11 Bacterial mRNA This figure compares a bacterial mRNA with the region of DNA from which it was transcribed. RNA polymerase recognizes, but does not transcribe, promoter sequences in the DNA. Initiation of RNA synthesis occurs five to eight bases downstream from the promoter. Ribosome recognition sites are located in the 5¿ mRNA upstream leader sequences. Protein-coding sequences begin at a start codon, which follows the leader sequences, and end at a downstream stop codon near the 3¿ end of the mRNA molecule. Downstream trailing sequences, which vary in length, follow the polypeptide-coding sequences.

KEY POINT

After transcription in eukaryotes, pre-mRNA undergoes extensive modification to produce mature, functional mRNA.

1st exon

Promoter

1st intron

2nd 2nd exon intron

3rd exon

mRNA termination sequence

Template DNA strand Transcription, capping of 5′ end

7-methylguanosine cap G– P – P – P

5′ end Stop codon

Start codon 1

Formation of pre-mRNA 1st intron

Small nuclear ribonucleoprotein complex

2nd intron –AAA... Poly-A tail 3′ end

G– P – P – P

5′ end 2

Processing of pre-mRNA (addition of poly-A tail and removal of introns)

1st exon

2nd exon

3rd exon –AAA... Poly-A tail 3′ end

G– P – P – P

5′ end 3

Protein-coding region

Mature mRNA in nucleus

Nuclear pore

Nuclear envelope

Cytosol

Transport through nuclear envelope to cytosol –AAA... Poly-A tail 3′ end Stop codon

G– P – P – P

5′ end 4

Start codon

Mature mRNA in cytosol

Protein-coding region

FIGURE 13-12 Animated Posttranscriptional modification of eukaryotic messenger RNA

the molecule persists in the cytosol), and facilitates initiation of translation by helping ribosomes recognize the mRNA. Both noncoding and coding sequences are transcribed from eukaryotic genes One of the greatest surprises in the history of molecular biology was the finding that most eukaryotic genes have interrupted coding sequences: long sequences of bases within the protein-coding sequences of the gene do not code for amino acids in the final polypeptide product. The noncoding regions within the gene are called

introns (intervening sequences), as opposed to exons (expressed sequences), which are parts of the protein-coding sequence. A typical eukaryotic gene may have multiple exons and introns, although the number varies. For example, the human b-globin gene, which produces one component of hemoglobin, contains 2 introns and 3 exons; the human gene for albumin contains 14 introns and 15 exons; and the human gene for titin, the largest protein known, found in muscle cells, contains 233 introns and 234 exons. In many cases, the combined lengths of the intron sequences are much greater than the combined lengths of the exons. For instance,

the ovalbumin gene contains about 7700 base pairs, whereas the total of all the exon sequences is only 1859 base pairs. When a gene that contains introns is transcribed, the entire gene is copied as a large RNA transcript, the pre-mRNA. A premRNA molecule contains both intron and exon sequences. (Note that intron and exon refer to corresponding nucleotide sequences in both DNA and RNA.) For the pre-mRNA to be made into a functional message, it must be capped and have a poly-A tail added, and the introns must be removed and the exons spliced together to form a continuous protein-coding message. Splicing itself occurs by several different mechanisms, depending on which type of RNA is involved. In many instances, splicing involves the association of several small nuclear ribonucleoprotein complexes (snRNPs, pronounced “snurps”) to form a large ribonucleoprotein complex called a spliceosome. Spliceosomes, which are similar in size to ribosomes, catalyze the reactions that remove introns from pre-mRNA. In other instances the RNA within the intron acts as an RNA catalyst, or ribozyme, splicing itself without the use of a spliceosome or protein enzymes. Posttranscriptional modification in eukaryotes is summarized as follows:

13.4 TRANSLATION ■ ■ LEARNING OBJECTIVES 8 Identify the features of tRNA that are important in decoding genetic information and converting it into “protein language.”

9 Explain how ribosomes function in polypeptide synthesis. 10 Describe the processes of initiation, elongation, and termination in polypeptide synthesis.

Translation adds another level of complexity to the process of information transfer because it involves conversion of the triplet nucleic acid code to the 20–amino acid alphabet of polypeptides. Translation requires the coordinated functioning of more than 100 kinds of macromolecules, including the protein and RNA components of the ribosomes, mRNA, and amino acids linked to tRNAs. Amino acids are joined by peptide bonds to form proteins. (See the formation of a peptide bond in Figure 3-18.) This joining links the amino and carboxyl groups of adjacent amino acids. The translation process ensures both that peptide bonds form and that the amino acids link in the correct sequence specified by the codons in the mRNA.

pre-mRNA containing introns and exons ¡ 5¿ end of pre-mRNA capped with modified nucleotide ¡ poly-A tail added to 3¿ end ¡ introns removed and exons spliced together ¡ mature mRNA transported into cytosol ¡ translation at ribosome

An amino acid is attached to tRNA before incorporation into a polypeptide

Following pre-mRNA processing, the mature mRNA is transported through a nuclear pore into the cytosol to be translated by a ribosome.

How do the amino acids align in the proper sequence so they can link? Crick proposed that a molecule was needed to bridge the gap between mRNA and proteins. This molecule was determined to be tRNA. DNA contains genes that are transcribed to form the tRNAs. Each kind of tRNA molecule binds to a specific amino acid. Amino acids are covalently linked to their respective tRNA molecules by enzymes, called aminoacyl–tRNA synthetases, which use ATP as an energy source (FIG. 13-13). The resulting complexes, called

Review ■

■

What features do mature eukaryotic mRNA molecules have that bacterial mRNAs lack?

■

In what ways are DNA polymerase and RNA polymerase similar? How do they differ? A certain template DNA strand has the following nucleotide sequence: 3¿¬TACTGCATAATGATT¬5¿ What would be the sequence of codons in the mRNA transcribed from this strand? What would be the nucleotide sequence of the complementary nontemplate DNA strand?

H H

N O

3′ H H

N

C

5′

Phenylalanine O C

O

ATP

+ H

A A G

O

AMP+ P P

Aminoacyl–tRNA synthetase

A A G

Anticodon Amino acid

tRNA

FIGURE 13-13 Linkage of a specific amino acid to the appropriate tRNA Amino acids are covalently linked to their respective tRNA molecules by aminoacyl–tRNA synthetases, which use ATP as an energy source.

Aminoacyl–tRNA

aminoacyl–tRNAs, bind to the mRNA coding sequence to align the amino acids in the correct order to form the polypeptide chain. The tRNAs are polynucleotide chains 70 to 80 nucleotides long, each with several unique base sequences as well as some sequences that are common to all. Although considerably smaller than mRNA or rRNA molecules, tRNA molecules have an intricate structure. A tRNA molecule has several properties essential to its function: (1) it has an anticodon, a complementary binding sequence for the correct mRNA codon; (2) it is recognized by an aminoacyl–tRNA synthetase that adds the correct amino acid; (3) it has an attachment site for the specific amino acid specified by the anticodon (the site lies about 180 degrees from the anticodon); and (4) it is recognized by ribosomes. Complementary base pairing within each tRNA molecule causes it to be doubled back and folded. Three or more loops of unpaired nucleotides are formed, one of which contains the anticodon. The amino acid–binding site is at the 3¿ end of the molecule. The carboxyl group of the amino acid is covalently bound to the exposed 3¿ hydroxyl group of the terminal nucleotide, leaving the amino group of the amino acid free to participate in peptide bond formation. The pattern of folding in tRNAs keeps a constant distance between the anticodon and amino acid, allowing precise positioning of the amino acids during translation.

The components of the translational machinery come together at the ribosomes The importance of ribosomes and of protein synthesis in cell metabolism is exemplified by a rapidly growing E. coli cell, which contains some 15,000 ribosomes—nearly one-third of the total mass of the cell. Although bacterial and eukaryotic ribosomes are not identical, ribosomes from all organisms share many fundamental features and basically work in the same way. Ribosomes consist of two subunits, both made up of protein (about 40% by weight) and rRNA (about 60% by weight). Unlike mRNA and tRNA, rRNA has catalytic functions and does not transfer information. The ribosomal proteins do not appear to be catalytic but instead contribute to the overall structure and stability of the ribosome. Researchers have isolated each ribosomal subunit intact in the laboratory, then separated each into its RNA and protein constituents. In bacteria, the smaller of these subunits contains 21 proteins and one rRNA molecule, and the larger subunit contains 34 proteins and two rRNA molecules. Under certain conditions researchers can reassemble each subunit into a functional form by adding each component in its correct order. Using this approach, together with sophisticated electron microscopic studies, researchers have determined the 3-D structure of the ribosome (FIG. 13-14a), as well as the way it is assembled in the living cell. The large subunit contains a depression on one surface into which the small subunit fits. During translation, the mRNA fits in a groove between the contact surfaces of the two subunits. The ribosome has four binding sites, one for mRNA and three for tRNAs. Thus, the ribosome holds the mRNA template, the aminoacyl–tRNA molecules, and the growing polypeptide chain in the correct orientation so the genetic code can be read and the next peptide bond formed. Transfer RNA molecules attach to three depressions on the ribosome, the A, P, and E binding sites

Front view

Large subunit

P A E P A site site

Large ribosomal subunit Ribosome

E site

P site

A site

Small subunit mRNA binding site

(a) A ribosome consists of two subunits, one large and one small. This model of a ribosome is based on a 3-D reconstruction of electron microscopic images.

Small ribosomal subunit

(b) The mRNA passes through a groove between the two ribosomal subunits. Each ribosome contains three binding sites for tRNA molecules.

FIGURE 13-14 Animated Ribosome structure

(FIG. 13-14b). The P site, or peptidyl site, is so named because the tRNA holding the growing polypeptide chain occupies the P site. The A site is named the aminoacyl site because the aminoacyl–tRNA delivering the next amino acid in the sequence binds at this location. The E site (for exit) is where tRNAs that have delivered their amino acids to the growing polypeptide chain exit the ribosome.

Translation begins with the formation of an initiation complex The process of protein synthesis has three distinct stages: initiation, repeating cycles of elongation, and termination. Initiation of protein synthesis is essentially the same in all organisms. Here we describe initiation in bacteria and then briefly discuss some differences between bacterial and eukaryotic initiation. The initiation of translation uses proteins called initiation factors, which become attached to the small ribosomal subunit. In bacteria, three different initiation factors attach to the small subunit, which then binds to the mRNA molecule in the region of the AUG start codon (FIG. 13-15). A leader sequence upstream (toward the 5¿ end) of the AUG helps the ribosome identify the AUG sequence that signals the beginning of the mRNA code for a protein. The tRNA that bears the first amino acid of the polypeptide is the initiator tRNA. The amino acid methionine is bound to the initiator tRNA, and as a result, the first amino acid on new poly-

KEY POINT

Initiation in bacteria involves a ribosome, an mRNA, an initiator tRNA to which formylated methionine (fMet) is bound, and several protein initiation factors.

3′

Leader sequence

Small ribosomal subunit 1

E site

5′

3′ mRNA

5′

Initiator tRNA

5′

P site

fMet

fMet

3′

Large ribosomal subunit A site

5′

3′

Initiation factor Start codon

The small ribosomal subunit binds to the mRNA at the AUG start codon. The leader sequence upstream of the AUG sequence helps the ribosome identify the AUG sequence.

Initiation complex 2

The initiator tRNA binds to the start codon, and one of the initiation factors is released.

3

When the large ribosomal subunit binds to the small subunit and the remaining initiation factors are released, the initiation complex is complete.

FIGURE 13-15 The initiation of translation in bacteria

peptides is methionine. (Quite often, the methionine is removed later.) In bacteria, the initiator methionine is modified by the addition of a one-carbon group derived from formic acid and is designated fMet for N-formylmethionine. The fMet–initiator tRNA, which has the anticodon 3¿¬UAC¬5¿, binds to the AUG start codon, releasing one of the initiation factors in the process. The initiation complex is complete when the large ribosomal subunit binds to the small subunit and the remaining initiation factors are released. In eukaryotes, initiation of translation differs in three respects. First, the methionine of the initiator tRNA is unmodified. Second, instead of a leader sequence to help identify the start codon on mRNA, the start codon is embedded within a short sequence (such as ACCAUGG) that indicates the site (the underlined AUG) of translation initiation. Third, the initiation complex, which contains perhaps 10 protein factors, is different from that of bacteria.

During elongation, amino acids are added to the growing polypeptide chain FIGURE 13-16 outlines the four steps in a cycle of elongation, the stage of translation in which amino acids are added one by one to the growing polypeptide chain. Elongation is essentially the same in bacteria and eukaryotes. The appropriate aminoacyl–tRNA recognizes the codon in the A site and binds there by base pairing of its anticodon with the complementary mRNA codon. This binding step requires several proteins called elongation factors. It also requires energy from guanosine triphosphate (GTP), an energy transfer molecule similar to ATP. The amino group of the amino acid at the A site is now aligned with the carboxyl group of the preceding amino acid attached to

the growing polypeptide chain at the P site. A peptide bond forms between the amino group of the new amino acid and the carboxyl group of the preceding amino acid. In this way, the polypeptide is released from tRNA at the P site and attaches to the aminoacyl– tRNA at the A site. This reaction is spontaneous (it does not require additional energy) because ATP transferred energy during formation of the aminoacyl–tRNA. Peptide bond formation does, however, require the enzyme peptidyl transferase. Remarkably, this enzyme is not a protein but a ribozyme that is an RNA component of the large ribosomal subunit. Two biochemists, American Thomas Cech (University of Colorado) and Canadian Sidney Altman (Yale University), received a Nobel Prize in Chemistry in 1989 for their independent discovery of ribozymes. Recall from Chapter 3 that polypeptide chains have direction, or polarity. The amino acid on one end of the polypeptide chain has a free amino group (the amino end), and the amino acid at the other end has a free carboxyl group (the carboxyl end). Protein synthesis always proceeds from the amino end to the carboxyl end of the growing polypeptide chain: 5′____________________ 3′

mRNA

Direction of translation

H

O N

H

C

polypeptide

OH

In translocation, the ribosome moves down the mRNA by one codon. As a result, the mRNA codon specifying the next amino acid in the polypeptide chain becomes positioned in the unoccupied A site. The translocation process requires energy, which again GTP supplies. The uncharged tRNA—that is, the tRNA without an attached amino acid—is moved from the P site

KEY POINT

Each repetition of the elongation process adds one amino acid to the growing polypeptide chain.

tRNA with an amino acid Amino acids

Amino acids GDP GTP

E

P

A

E

P

A

Aminoacyl–tRNA binds to codon in A site 5′

1

3′

mRNA

3′

5′

The polypeptide chain is covalently bonded to the tRNA that carries the amino acid most recently added to the chain. This tRNA is in the P site of the ribosome.

2

An aminoacyl–tRNA binds to the A site by complementary base pairing between the tRNA’s anticodon and the mRNA’s codon.

Ribosome ready to accept another aminoacyl–tRNA

Peptide bond formation

Amino end of polypeptide

New peptide bond

E

P

Translocation toward 3′ end of mRNA

A

E

P

A

GTP GDP 5′

4

3′

In the translocation step, the ribosome moves one codon toward the 3′ end of mRNA. As a result, the growing polypeptide chain is transferred to the P site. The uncharged tRNA in the E site exits the ribosome.

5′

3′

3

The growing polypeptide chain detaches from the tRNA molecule in the P site and becomes attached by a peptide bond to the amino acid linked to the tRNA at the A site.

FIGURE 13-16 An elongation cycle in translation This illustration begins after a short chain of amino acids has formed.

to the E site. From there it exits the ribosome and joins the cytosolic pool of tRNAs. Because translocation involves movement of the ribosome in the 3¿ direction along the mRNA molecule, translation always proceeds in the 5¿ ¡ 3¿ direction. Each peptide bond forms in a fraction of a second; by repeating the elongation cycle, an averagesized protein of about 360 amino acids is assembled by a bacterium in approximately 18 seconds and by a eukaryotic cell in a little more than 1 minute.

One of three stop codons signals the termination of translation Termination is the final stage of translation. In termination, the synthesis of the polypeptide chain is terminated by a release factor, a protein that recognizes the stop codon at the end of the coding sequence. (Because no tRNA molecule binds to a stop codon, the codon is available for binding by the release factor.) When the release factor binds to the A site, the bond between the tRNA in

KEY POINT

A stop signal terminates polypeptide synthesis, because no tRNA molecules exist with an anticodon complementary to a stop codon. Large ribosomal subunit Polypeptide chain is released

Release factor E

5′

P

A

E

3′

mRNA

5′

1

When the ribosome reaches a stop codon, the A site binds to a protein release factor.

A

Release factor

3′ Stop codon (UAA, UAG, or UGA)

2

P

P

mRNA

Stop codon (UAA, UAG, or UGA)

E

5′

The release factor hydrolyzes the bond between the polypeptide chain and the tRNA, causing the release of the polypeptide chain from the tRNA molecule in the P site.

mRNA Small ribosomal subunit tRNA 3

3′

The remaining parts of the translation complex dissociate.

FIGURE 13-17 The termination of translation

the P site and the last amino acid of the polypeptide chain breaks (FIG. 13-17). This hydrolysis reaction frees the newly synthesized polypeptide and also separates the translation complex into its parts: the mRNA molecule, the release factor, the tRNA molecule in the P site, and the large and small ribosomal subunits. The two ribosomal subunits can be used to form a new initiation complex with another mRNA molecule. Each mRNA is translated only a limited number of times before it is destroyed. Specialized ribosome-associated proteins, called molecular chaperones, assist in the folding of the newly synthesized polypeptide chain into its three-dimensional active shape (see Chapter 3). The sequence of amino acids in the polypeptide chain dictates the shape that it ultimately forms. However, without the molecular chaperone’s help, interactions among the various regions of the amino acid chain might prevent the proper folding process from occurring.

Review ■

■

■

What are ribosomes made of? Do ribosomes carry information to specify the amino acid sequence of proteins? What happens in each stage of polypeptide synthesis: initiation, elongation, and termination? A certain mRNA strand has the following nucleotide sequence: 5¿¬AUG¬ACG¬UAU¬AAC¬UUU¬3¿ What is the anticodon for each codon? What is the amino acid sequence of the polypeptide? (Use Fig. 13-5 to help answer this question.)

13.5 VARIATIONS IN GENE EXPRESSION ■ ■ LEARNING OBJECTIVES 11 Describe a polyribosome in bacterial cells. 12 Briefly discuss RNA interference. 13 Describe retroviruses and the enzyme reverse transcriptase.

You have seen that the basic mechanisms of transcription and translation are quite similar in all organisms. Some significant differences exist between bacteria and eukaryotes (FIG. 13-18). Some of these differences result from bacterial versus eukaryotic cell structure. Bacteria lack a nucleus, whereas eukaryotic chromosomes are confined to the cell nucleus and protein synthesis takes place in the cytosol. In eukaryotes, mRNA must move from the nucleus (the site of transcription) to the cytosol (the site of translation, or polypeptide synthesis). Also, as we discussed earlier in the chapter, extensive mRNA processing occurs in eukaryotic cells before translation.

Transcription and translation are coupled in bacteria In E. coli and other bacteria, translation begins very soon after transcription, and several ribosomes are attached to the same mRNA (FIG. 13-19). The end of the mRNA molecule synthesized first during transcription is also the first to be translated to form a polypeptide.

Eukaryotic cell

In the early 1980s, American Walter Gilbert of Harvard University proBacterial cell posed that exons are nucleNucleus otide sequences that code for protein domains, regions DNA of protein tertiary structure that may have important Transcription DNA functions. For example, the Pre-mRNA active site of an enzyme may compose one domain. A difRNA processing Transcription ferent domain may enable mRNA that enzyme to bind to a mRNA particular cell structure, and yet another may be a site involved in allosteric regulation Translation Translation (see Chapter 7). Analyses of the DNA and amino acid sePolypeptide Polypeptide quences of many eukaryotic Ribosome Ribosome genes have shown that most exons are too small to code for an entire protein domain, although a block of several exons can code for a domain. Gilbert further postu(a) In a bacterial cell, mRNA is (b) In a eukaryotic cell, RNA processing occurs in the lated that new proteins with immediately ready for translation nucleus before the mRNA exits the nucleus for translation. new functions emerge rapidly by ribosomes. when genetic recombination produces novel combinations FIGURE 13-18 Summary: the flow of genetic information in bacteria and eukaryotes of exons that code for different proteins. This hypothesis has become known as evolution by exon shuffling. It has been supported Ribosomes bind to the 5¿ end of the growing mRNA and initiate by examples such as the low-density lipoprotein (LDL) receptor translation long before the message is fully synthesized. As many protein, a protein found on the surface of human cells that binds to as 15 ribosomes may bind to a single mRNA. An mRNA molecule cholesterol transport molecules (see Chapter 5). The LDL receptor that is bound to clusters of ribosomes constitutes a polyribosome, protein has domains that are related to parts of other proteins with or polysome. Polyribosomes also occur in eukaryotic cells, but not totally different functions. However, many other genes and their while transcription is still occurring. corresponding proteins show no evidence of exon shuffling. Although many polypeptide chains can be actively syntheAfter introns were discovered in a few genes in archaea and sized on a single mRNA at any one time, the half-life of mRNA bacteria, some biologists hypothesized that introns are relics from molecules in bacterial cells is only about 2 minutes. (Half-life is the the ancient ancestors of all organisms living today. Through biltime it takes for half the molecules to be degraded.) Usually, degralions of years of evolution, the pressure in prokaryotic organisms, dation of the 5¿ end of the mRNA begins even before the first polywhich are unicellular, to have a streamlined set of genes may have peptide is complete. Once the ribosome recognition sequences at caused them to lose most of their introns. the 5¿ end of the mRNA have been degraded, no more ribosomes Regardless of how split genes originated, intron excision proattach and initiate protein synthesis. vides one of many ways in which present-day eukaryotes regulate expression of their genes (see Chapter 14). This opportunity for Biologists debate the evolution control, together with the fact that eukaryotic RNAs are far more of eukaryotic gene structure stable than those of bacteria, may balance the energy cost of mainThe reason for the intricate structure of eukaryotic genes is a mattaining a large load of noncoding DNA. ter of ongoing discussion among molecular biologists. Why do introns occur in most eukaryotic nuclear genes but not in the genes Several kinds of eukaryotic RNA of most bacteria (or of mitochondria and chloroplasts)? How did have a role in gene expression this remarkable genetic system with interrupted coding sequences In addition to mRNA, rRNA, and tRNA, eukaryotic cells contain (“split genes”) evolve, and why has it survived? It seems incredseveral other kinds of RNA that are an essential part of the comible that as much as 75% of the original transcript of a eukaryotic plex system that results in protein synthesis (TABLE 13-1). Small nuclear gene must be removed to make a working message.

Inactive DNA segment

Courtesy of Dr. Barbara Hamkalo, University of California at Irvine

RNA polymerase Active DNA segment

Direction of RNA synthesis

Active DNA segment

Ribosomes Polyribosome Direction of protein synthesis 0.5 μm mRNA

(a) This TEM shows two strands of E. coli DNA, one inactive and the other actively producing mRNA. Protein synthesis begins while the mRNA is being completed.

(b) A sequence (left to right ) of coupled transcription and translation. Note that several ribosomes translate each mRNA molecule simultaneously.

FIGURE 13-19 Coupled transcription and translation in bacteria

malian cells, about 48% of the pre-rRNA molecule is removed durnuclear RNA (snRNA) molecules bind to specific proteins to ing the cleavage steps. Pre-rRNA may have methyl groups added to form a small nuclear ribonucleoprotein complex (snRNP), which regions that are to be saved during processing. in turn combines with other snRNPs to form a spliceosome. (ReThe discovery that RNA has the ability to regulate gene exprescall from the discussion earlier in the chapter that a spliceosome sion made big news in biology, both because this phenomenon, catalyzes intron removal.) known as RNA interference (RNAi), is found in many organisms Some polypeptides enter the endoplasmic reticulum (ER) and because it has great potential as a research and medical tool as they are being synthesized. Signal-recognition particle RNA (see Chapters 15 and 17). In RNAi, certain small RNA molecules (SRP RNA), in combination with proteins, directs the ribosome– mRNA–polypeptide complex to the rough ER shortly after translation begins. TABLE 13-1 Selected Kinds of RNA in Eukaryotic Cells Translation continues after the complex Kind of RNA Function binds to an integral protein in the ER membrane, and the newly synthesized Messenger RNA (mRNA) Specifies the amino acid sequence of a protein protein is then distributed to its destinaTransfer RNA (tRNA) Binds to specific amino acid and serves as adapter molecule when tion within the cell or outside the cell (for amino acids are incorporated into growing polypeptide chain example, as a secretion). Ribosomal RNA (rRNA) Has both structural and catalytic (ribozyme) roles in ribosome A special group of RNA molecules, Small nuclear RNA (snRNA) Involved in intron removal and regulation of transcription; part of spliceosome particles known as small nucleolar RNAs (snoSignal-recognition particle Helps direct the ribosome–mRNA–polypeptide complex to the RNAs), processes pre-rRNA molecules RNA (SRP RNA) rough ER shortly after translation begins in the nucleolus during ribosome subSmall nucleolar RNA (snoRNA) Processes pre-rRNA in nucleolus during formation of ribosome unit formation. Small nucleolar RNAs subunits bind to complementary regions of preSmall interfering RNA (siRNA) Controls transposons and fights viral infections rRNA and target sites for cleavage or MicroRNA (miRNA) Controls expression of genes involved in growth and developmethylation. A pre-rRNA molecule is ment by inhibiting the translation of mRNA; involved in several diseases, including prevention of cancer larger than the combined size of the three rRNA molecules made from it. In mam-

The definition of a gene has evolved as biologists have learned more about genes At the beginning of this chapter, we traced the development of ideas regarding the nature of the gene. For a time scientists found it useful to define a gene as “a sequence of nucleotides that codes for one polypeptide chain.” However, as we have learned more about the human genome, research teams have determined that DNA coding for polypeptides makes up about 2% of our genome, whereas about 80% of our genome is expressed; many genes are transcribed to produce the various kinds of RNA molecules (see Table 13-1). Further, as you will see in Chapter 14, studies have shown that in eukaryotic cells, a single gene may produce more than one polypeptide chain by modifications in the way the mRNA is processed. It is perhaps most useful to define a gene in terms of its product: a gene is a DNA nucleotide sequence that carries the information needed to produce a specific RNA or polypeptide product.

The usual direction of information flow has exceptions For several decades, a central premise of molecular biology was that genetic information always flows from DNA to RNA to protein. Through his studies of viruses, U.S. biologist Howard Temin discovered an important exception to this rule in 1964. Although viruses are not cellular organisms, they contain a single type of nucleic acid and reproduce in a host cell. Temin was studying unusual, cancer-causing tumor viruses that have RNA, rather than DNA, as their genetic material. He found that infection of a host cell by one of these viruses is blocked by inhibitors of DNA synthesis and also by inhibitors of transcription. These findings suggested

Javier F. Palatnik and Detlef Weigel

interfere with the expression of genes or their RNA transcripts. RNA interference involves small interfering RNAs, microRNAs, and a few other kinds of short RNA molecules. Small interfering RNAs (siRNAs) are double-stranded molecules that are 20 to 25 nucleotides in length. These RNA molecules help control damage from transposons (discussed later in the chapter) and viral infections; they also regulate gene expression of protein-coding genes. MicroRNAs (miRNAs) are single-stranded RNA molecules 20 to 25 nucleotides long that inhibit the translation of mRNAs involved in many biological processes, such as growth and development (FIG. 13-20). MicroRNAs are transcribed from genes and then shortened before they are combined with proteins to form a complex that inhibits the expression of mRNA molecules with sequences complementary to the miRNA. More than 200 distinct miRNA genes have been identified in the human genome, but the specifics of how they regulate gene expression by degrading mRNAs and/or preventing mRNAs from being translated into polypeptides are not yet clear. Biologists now recognize that miRNAs have an important role in preventing cancer: cancer cells have measurably lower levels of miRNAs than noncancer cells. A breakdown in miRNA function has also been implicated in heart disease and possibly Parkinson’s disease.

FIGURE 13-20 The effect of an miRNA molecule on plant growth A normal Arabidopsis plant (top view, leaf, and side view of seedling) is shown on the left and serves as a control. Biologists modified an miRNA molecule in the plant on the right, and as a result, it exhibits severe developmental abnormalities, such as curly leaves.

that DNA synthesis and transcription are required for RNA tumor viruses to multiply and that there must be a way for information to flow in the “reverse” direction—from RNA to DNA. Temin proposed that a DNA provirus forms as an intermediary in the replication of RNA tumor viruses. This hypothesis required a new kind of enzyme that would synthesize DNA using RNA as a template. In 1970, Temin and U.S. biologist David Baltimore discovered just such an enzyme, and they shared the Nobel Prize in Physiology or Medicine in 1975 for their discovery. This RNA-directed DNA polymerase, also known as reverse transcriptase, is found in all RNA tumor viruses. (Some RNA viruses that do not produce tumors, however, replicate without using a DNA intermediate.) FIGURE 13-21 shows the steps of RNA tumor virus reproduction. Because they reverse the usual direction of information flow, viruses that require reverse transcriptase are called retroviruses. HIV-1, the virus that causes AIDS, is the most widely known retrovirus. As you will see in Chapter 15, the reverse transcriptase enzyme has become an important research tool for molecular biologists.

Review ■ ■ ■

What is RNA interference? What is a gene? How do retroviruses use the enzyme reverse transcriptase?

13.6 MUTATIONS ■ ■ LEARNING OBJECTIVE 14 Give examples of the different classes of mutations that affect the base sequence of DNA and explain the effects that each has on the polypeptide produced.

One of the first major discoveries about genes was that they undergo mutations, changes in the nucleotide sequence of the DNA. However, the overall mutation rate is much higher than the frequency of damage to DNA because all organisms have systems of enzymes that repair certain kinds of DNA damage.

Chromosome DNA in nucleus of host cell

DNA provirus

RNA/DNA hybrid Viral RNA

Provirus inserted into chromosome DNA

Provirus DNA transcribed

DNA replication

Viral mRNA

Digestion of RNA strand

Viral RNA

Reverse transcription

Viral proteins RNA virus

RNA virus

(a) After an RNA tumor virus enters the host cell, the viral enzyme reverse transcriptase synthesizes a DNA strand that is complementary to the viral RNA. Next, the RNA strand is degraded and a complementary DNA strand is synthesized, thus completing the double-stranded DNA provirus, which is then integrated into the host cell’s DNA.

(b) The provirus DNA is transcribed, and the resulting viral mRNA is translated to form viral proteins. Additional viral RNA molecules are produced and then incorporated into mature viral particles enclosed by protein coats.

FIGURE 13-21 The infection cycle of an RNA tumor virus

If a DNA sequence has been changed and not corrected, DNA replication copies the altered sequence just as it would copy a normal sequence, making the mutation stable over an indefinite number of generations. In most cases, the mutant allele has no greater tendency than the original allele to mutate again. Although most uncorrected mutations are either silent (and have no discernible effect) or harmful (such as those that cause inherited diseases and cancer), a few are useful. Because they can be passed to the next generation in cell lines that give rise to gametes, mutations are of vital importance in evolution. Mutations provide the variation among individuals that evolutionary forces act on. Mutations are also useful in research (recall Beadle and Tatum’s work with mutations) because they provide the diversity of genetic material that enables researchers to study inheritance and the molecular nature of genes. Scientists now determine where a particular mutation occurs in a gene by using recombinant DNA methods to isolate the gene and determine its sequence of bases (see Chapter 15).

Base-pair substitution mutations result from the replacement of one base pair by another Mutations can alter genes in several ways. The simplest type of mutation, called a base-pair substitution, involves a change in only one pair of nucleotides. Often these mutations result from errors in base pairing during the replication process. For example, a GC, CG, or TA pair might replace an AT base pair. Such a mutation may cause the altered DNA to be transcribed as an altered mRNA. The altered mRNA may then be translated into a polypeptide chain with only one amino acid different from the normal sequence.

Silent mutations are base-pair substitutions that have no discernible effect—as, for example, a mutation in a protein-coding gene that does not alter the amino acid sequence (FIGS. 13-22a, b). Base-pair substitutions that result in replacement of one amino acid by another are sometimes called missense mutations (FIG. 13-22c). Missense mutations have a wide range of effects. If the amino acid substitution occurs at or near the active site of an enzyme, the activity of the altered protein may decrease or even be destroyed. Some missense mutations involve a change in an amino acid that is not part of the active site. Others may result in the substitution of a closely related amino acid (one with very similar chemical characteristics). Such mutations have no effect on the function of a gene product and may be undetectable. Because these mutations occur relatively frequently, the true number of mutations in an organism is much greater than is actually observed. Nonsense mutations are base-pair substitutions that convert an amino acid–specifying codon to a stop codon (FIG. 13-22d). A nonsense mutation usually destroys the function of the gene product; in the case of a protein-specifying gene, the part of the polypeptide chain that follows the mutant stop codon is missing.

Frameshift mutations result from the insertion or deletion of base pairs In frameshift mutations, one or two nucleotide pairs are inserted into or deleted from the molecule, altering the reading frame. As a result, codons downstream of the insertion or deletion site specify an entirely new sequence of amino acids. Depending on where the insertion or deletion occurs in the gene, different effects are generated. Frameshift mutations may produce a stop

(a) Normal DNA sequence

(b) Base-pair substitution (silent mutation)

DNA template strand 3′

TAC

TGA

TCT

TTA

ACC

CTA

5′

3′

TAC

TGA

TCC

TTA

ACC

CTA

5′

5′

ATG

ACT

AGA

AAT

TGG

GAT

3′

5′

ATG

ACT

AGG

AAT

TGG

GAT

3′

AAU

UGG

GAU

3′

5′

AUG

ACU

AGG

AAU

UGG

GAU

3′

Asn

Trp

Asp

Met

Thr

Arg

Asn

Trp

Asp

DNA (normal sequence)

5′

AUG

ACU

AGA

mRNA (normal sequence)

Met

Thr

Arg

Polypeptide (normal sequence)

(c) Base-pair substitution (missense)

(d) Base-pair substitution (nonsense)

3′

TAC

TGA

TGT

TTA

ACC

CTA

5′

3′

TAC

TGA

TCT

TTA

ATC

CTA

5′

5′

ATG

ACT

ACA

AAT

TGG

GAT

3′

5′

ATG

ACT

AGA

AAT

TAG

GAT

3′

5′

AUG

ACU

ACA

AAU

UGG

GAU

3′

5′

AUG

ACU

AGA

AAU

UAG

GAU

3′

Met

Thr

Thr

Asn

Trp

Asp

Met

Thr

Arg

Asn

(Stop)

(e) Frameshift (nonsense)

(f) Frameshift (altered amino acid sequence)

T G missing

T missing

3′

TAC

ATC

TTT

AAC

CCT

AGG

5′

3′

TAC

TGA

CTT

TAA

CCC

TAG

5′

5′

ATG

TAG

AAA

TTG

GGA

TCC

3′

5′

ATG

ACT

GAA

ATT

GGG

ATC

3′

3′

A C missing 5′

AUG

UAG

Met

(Stop)

A missing AAA

UUG

GGA

UCC

3′

5′

AUG

ACU

GAA

AUU

GGG

AUC

Met

Thr

Glu

Ile

Gly

Ile

▲

FIGURE 13-22 Animated Types of mutations

Compare the normal DNA sequence in part (a) with the mutations in parts (b) to (e).

codon within a short distance of the mutation (FIG. 13-22e). This codon terminates the already altered polypeptide chain. In addition to producing a stop codon, frameshift mutations can result in an altered amino acid sequence immediately after the change (FIG. 13-22f). A frameshift mutation in a gene specifying an enzyme usually results in a complete loss of enzyme activity.

Some mutations involve mobile genetic elements One type of mutation is caused by DNA sequences that “jump” into the middle of a gene. These movable sequences of DNA are known as mobile genetic elements, or transposons. Mobile genetic elements not only disrupt the functions of some genes but also inactivate some previously active genes. Transposons were discovered in maize (corn) by the U.S. geneticist Barbara McClintock in the 1950s (FIG. 13-23). She observed that certain genes appeared to be turned off and on spontaneously. She deduced that the mechanism involved a segment of DNA that moved from one region of a chromosome to another, where it would inactivate a gene. In recognition of her insightful findings, McClintock was awarded the Nobel Prize in Physiology or Medicine in 1983. Biologists did not appreciate the significance of transposons until the development of recombinant DNA methods and the discovery of transposons in a wide variety of organisms, including hu-

mans. We now know that transposons are segments of DNA that range from a few hundred to several thousand bases. In humans, almost half of our genome consists of transposons. Several kinds of mobile genetic elements have been identified. One of these, called a DNA transposon, moves genetic material from one site to another using a “cut-and-paste” method. DNA transposons are common in bacteria and animals, including humans. Many mobile genetic elements are retrotransposons, which replicate by forming an RNA intermediate. Reverse transcriptase converts them into their original DNA sequence before they jump into a gene. Many retrotransposons enlist a host cell’s gene expression machinery to produce reverse transcriptase encoded by the retrotransposon. Because retrotransposons use reverse transcriptase, some biologists hypothesize that certain retrotransposons may have evolved from retroviruses, or vice versa. Scientists think that human cells have fewer than 100 retrotransposons actively jumping around. Nevertheless, retrotransposons are extremely important agents of mutation and therefore increase an organism’s ability to evolve. By one estimate, retrotransposons are responsible for about 10% of naturally occurring mutations that cause noticeable phenotypic changes.

Mutations have various causes

Jerome Wexler/Visuals Unlimited

Most types of mutations occur infrequently but spontaneously from mistakes in DNA replication or defects in the mitotic or meiotic separation of chromosomes. Some regions of DNA, known as mutational hot spots, are much more likely than others to undergo mutation. An example is a short stretch of repeated nucleotides, which can cause DNA polymerase to “slip” while reading the template during replication. Mutations in certain genes increase the overall mutation rate. For example, a mutation in a gene coding for DNA polymerase may make DNA replication less precise, or a mutation in a gene coding for a repair enzyme may allow more mutations to arise as a result of unrepaired DNA damage. Not all mutations occur spontaneously. Mutagens, which cause many of the types of mutations discussed previously, include various types of radiation—such as X-rays, gamma rays, cosmic rays, and ultraviolet rays—and certain chemicals. Some chemical mutagens react with and modify bases in the DNA, leading to mistakes in complementary base pairing when the DNA molecule is replicated. Other chemical mutagens cause nucleotide pairs to be inserted into or deleted from the DNA molecule, changing the normal reading frame FIGURE 13-23 Transposons in maize during replication. The white lines and streaks in these kernels are caused by transposons that have moved from one Despite the presence of enzymes that relocation to another, turning off pigment production in certain cells. The number of cell divisions durpair damage to DNA, some new mutations do ing the development of the kernel since the transposon turned off the gene determines how much arise. In fact, each of us has some mutant alpigment is produced. Some kernels are colorless (unpigmented) because the pigment-producing genes were turned off very early in development. leles that neither of our parents had. Although

some of these mutations alter the phenotype, most are not noticeable because they are recessive. Mutations that occur in the cells of the body (somatic cells) are not passed on to offspring. However, these mutations are of concern because somatic mutations and cancer are closely related. Many mutagens are also carcinogens, agents that cause cancer.

■ ■

Review What are the main types of mutations? What effects does each type of mutation have on the polypeptide product produced?

■ ■

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

13.1 (page 283) 1 Summarize the early evidence indicating that most genes specify the structure of proteins. Garrod’s work on inborn errors of metabolism provided evidence that genes specify proteins. Garrod studied a rare genetic disease called alkaptonuria and hypothesized that people with alkaptonuria lack the enzyme that normally oxidizes homogentisic acid. 2 Describe Beadle and Tatum’s experiments with Neurospora. ■ Beadle and Tatum exposed Neurospora spores to X-rays or ultraviolet radiation to induce mutant strains, and then identified strains that carried a mutation preventing the fungus from producing a chemical essential for growth. Their work revealed that each mutant strain had a mutation in only one gene and that each gene affected only one enzyme. ■

13.2 (page 285) 3 Outline the flow of genetic information in cells, from DNA to RNA to polypeptide. The process by which information encoded in DNA specifies the sequences of amino acids in proteins involves transcription and translation. During transcription, an RNA molecule complementary to the template DNA strand is synthesized. Messenger RNA (mRNA) molecules contain information that specifies the amino acid sequences of polypeptide chains. ■ During translation, a polypeptide chain specified by the mRNA is synthesized. Each sequence of three nucleotide bases in the mRNA constitutes a codon, which specifies one amino acid in the polypeptide chain, or a start or stop signal. Translation requires tRNAs and cell machinery, including ribosomes. ■

enine, guanine, or cytosine), and three phosphates. Like DNA, RNA subunits are covalently joined by 5¿¬3¿ linkages to form an alternating sugar–phosphate backbone.

5 Explain why the genetic code is said to be redundant and virtually universal, and discuss how these features may reflect its evolutionary history. ■ The genetic code is read from mRNA as a series of codons that specify a single sequence of amino acids. The genetic code is redundant because some amino acids are specified by more than one codon. Only a few minor variations are exceptions to the genetic code found in all organisms, strongly suggesting that all organisms are descended from a common ancestor.

13.3 (page 288) 6 Compare the processes of transcription and DNA replication, identifying both similarities and differences. ■ RNA polymerases, involved in RNA synthesis, have many similarities to DNA polymerases involved in DNA replication. Both enzymes carry out synthesis in the 5¿ ¡ 3¿ direction. Both use nucleotides with three phosphate groups as substrates, removing two of the phosphates as the nucleotides are covalently linked to the 3¿ end of the newly synthesized strand. ■ Just as the paired strands of DNA are antiparallel, the template strand of the DNA and its complementary RNA strand are antiparallel. As a result, the DNA template strand is read in its 3¿ ¡ 5¿ direction as RNA is synthesized in its 5¿ ¡ 3¿ direction. ■ The same base-pairing rules are followed as in DNA replication, except that uracil is substituted for thymine. Watch transcription unfold by clicking on the figure in CengageNOW.

7 Compare bacterial and eukaryotic mRNAs, and explain the functional DNA Transcription mRNA Translation Polypeptide Ribosome

significance of their structural differences. ■ Eukaryotic genes and their mRNA molecules are more complicated than those of bacteria. After transcription, a 5¿ cap (a modified guanosine triphosphate) is added to the 5¿ end of a eukaryotic mRNA molecule. The molecule also has a poly-A tail of adeninecontaining nucleotides added at the 3¿ end. These modifications may protect eukaryotic mRNA molecules from degradation, giving them a longer lifespan than bacterial mRNA. ■ In many eukaryotic genes, the coding regions, called exons, are interrupted by noncoding regions, called introns. Both exons and introns are transcribed, but the introns are later removed from the original pre-mRNA, and the exons are spliced together to produce a continuous polypeptide-coding sequence.

13.4 (page 293) 8 Identify the features of tRNA that are important in decoding genetic 4 Compare the structures of DNA and RNA. ■

Like DNA, RNA is formed from nucleotide subunits. However, in RNA each subunit contains the sugar ribose, a base (uracil, ad-

information and converting it into “protein language.” ■ Transfer RNAs (tRNAs) are the “decoding” molecules in the translation process. Each tRNA molecule is specific for only one amino acid. One part of the tRNA molecule contains an anticodon that is

complementary to a codon of mRNA. Attached to one end of the tRNA molecule is the amino acid specified by the complementary mRNA codon.

9 Explain how ribosomes function in polypeptide synthesis. ■

Each ribosome is made of a large and a small subunit; each subunit contains ribosomal RNA (rRNA) and many proteins. Ribosomes couple the tRNAs to their proper codons on the mRNA, catalyze the formation of peptide bonds between amino acids, and translocate the mRNA so the next codon can be read. Learn more about the structure of ribosomes by clicking on the figure in CengageNOW.

10 Describe the processes of initiation, elongation, and termination in polypeptide synthesis. ■ Initiation is the first stage of translation. Initiation factors bind to the small ribosomal subunit, which then binds to mRNA in the region of AUG, the start codon. The initiator tRNA binds to the start codon, followed by binding of the large ribosomal subunit. ■ Elongation is a cyclic process in which amino acids are added one by one to the growing polypeptide chain. Elongation proceeds in the 5¿ ¡ 3¿ direction along the mRNA. ■ Termination, the final stage of translation, occurs when the ribosome reaches one of three stop codons. The A site binds to a release factor, which triggers the release of the completed polypeptide chain and dissociation of the translation complex. Learn more about protein synthesis by clicking on the figure in CengageNOW.

13.5 (page 297) 11 Describe a polyribosome in bacterial cells. ■

Unlike the processes in eukaryotic cells, transcription and translation are coupled in bacterial cells. Bacterial ribosomes bind to the

5¿ end of the growing mRNA and initiate translation before the message is fully synthesized. As many as 15 ribosomes may bind to a single mRNA, forming a polyribosome. 12 Briefly discuss RNA interference. ■ RNA interference (RNAi) occurs when small RNA molecules, such as small interfering RNAs (siRNAs) and microRNAs (miRNAs) interfere with the expression of genes or their RNA transcripts.

13 Describe retroviruses and the enzyme reverse transcriptase. ■

Retroviruses are viruses that synthesize DNA from an RNA template. In retroviruses the flow of genetic information is reversed by the enzyme reverse transcriptase. HIV-1, the virus that causes AIDS, is a retrovirus.

13.6 (page 300) 14 Give examples of the different classes of mutations that affect the base sequence of DNA and explain the effects that each has on the polypeptide produced. ■ A base-pair substitution is a mutation that may alter or destroy the function of a protein if a codon changes so that it specifies a different amino acid (a missense mutation) or becomes a stop codon (a nonsense mutation). ■ The insertion or deletion of one or two base pairs in a gene invariably destroys the function of that protein because it results in a frameshift mutation, which changes the codon sequences downstream from the mutation. ■ One type of mutation is caused by movable DNA sequences, known as transposons, that “jump” into the middle of a gene. Many transposons are retrotransposons, which replicate by forming an RNA intermediate; reverse transcriptase converts them to their original DNA sequence before they jump into a gene. Learn more about mutations by clicking on the figure in CengageNOW.

T E S T YO U R U N D E R S TA N D I N G 1. Beadle and Tatum (a) predicted that tRNA molecules would have anticodons (b) discovered the genetic disease alkaptonuria (c) showed that the genetic disease sickle cell anemia is caused by a change in a single amino acid in a hemoglobin polypeptide chain (d) worked out the genetic code (e) studied the relationship between genes and enzymes in Neurospora 2. What is the correct order of information flow in bacterial and eukaryotic cells? (a) DNA ¡ mRNA ¡ protein (b) protein ¡ mRNA ¡ DNA (c) DNA ¡ protein ¡ mRNA (d) protein ¡ DNA ¡ mRNA (e) mRNA ¡ protein ¡ DNA 3. During transcription, how many RNA nucleotide bases would usually be encoded by a sequence of 99 DNA nucleotide bases? (a) 297 (b) 99 (c) 33 (d) 11 (e) answer is impossible to determine with the information given 4. The genetic code is defined as a series of _____ in _____. (a) anticodons; tRNA (b) codons; DNA (c) anticodons; mRNA (d) codons; mRNA (e) codons and anticodons; rRNA 5. RNA differs from DNA in that the base _____ is substituted for _____. (a) adenine; uracil (b) uracil; thymine (c) guanine; uracil (d) cytosine; guanine (e) guanine; adenine

6. RNA grows in the _____ direction, as RNA polymerase moves along the template DNA strand in the _____ direction. (a) 5¿ ¡ 3¿; 3¿ ¡ 5¿ (b) 3¿ ¡ 5¿; 3¿ ¡ 5¿ (c) 5¿ ¡ 3¿; 5¿ ¡ 3¿ (d) 3¿ ¡ 3¿; 5¿ ¡ 5¿ (e) 5¿ ¡ 5¿; 3¿ ¡ 3¿ 7. Which of the following is/are not found in a bacterial mRNA molecule? (a) stop codon (b) upstream leader sequences (c) downstream trailing sequences (d) start codon (e) promoter sequences 8. Which of the following is/are typically removed from pre-mRNA during nuclear processing in eukaryotes? (a) upstream leader sequences (b) poly-A tail (c) introns (d) exons (e) all the preceding 9. The role of tRNA is to transport (a) amino acids to the ribosome (b) amino acids to the nucleus (c) initiation factors to the ribosome (d) mRNA to the ribosome (e) release factors to the ribosome 10. Suppose you mix the following components of protein synthesis in a test tube: amino acids from a rabbit; ribosomes from a dog; tRNAs from a mouse; mRNA from a chimpanzee; and necessary enzymes plus an energy source from a giraffe. If protein synthesis occurs, which animal’s protein will be made? (a) rabbit (b) dog (c) mouse (d) chimpanzee (e) giraffe

11. The _____ catalyzes the excision of introns from pre-mRNA. (a) ribosome (b) spliceosome (c) RNA polymerase enzyme (d) aminoacyl–tRNA synthetase enzyme (e) reverse transcriptase enzyme 12. A nonsense mutation (a) causes one amino acid to be substituted for another in a polypeptide chain (b) results from the deletion

of one or two bases, leading to a shift in the reading frame (c) results from the insertion of one or two bases, leading to a shift in the reading frame (d) results from the insertion of a transposon (e) usually results in the formation of an abnormally short polypeptide chain

CRITICAL THINKING 1. Compare and contrast the formation of mRNA in bacterial and eukaryotic cells. How do the differences affect the way in which each type of mRNA is translated? Does one system have any obvious advantage in terms of energy cost? Which system offers greater opportunities for control of gene expression? 2. Biologists hypothesize that transposons eventually lose the ability to replicate and therefore remain embedded in DNA without moving around. Based on what you have learned in this chapter, suggest a possible reason for this loss. 3. How was the genetic code deciphered? 4. Draw an arrow on this figure to show where reverse transcription, catalyzed by the enzyme reverse transcriptase, occurs. Also, show where the enzymes DNA polymerase, RNA polymerase, and peptidyl transferase are involved. DNA replication

Transcription DNA

Translation RNA

Polypeptide

5. ANALYZING DATA. You are repeating Beadle and Tatum’s classic experiment in biology lab. First, you expose Neurospora spores to X-rays, establish a culture from an irradiated spore, and then grow it on (1) a minimal medium plus vitamins, (2) a minimal medium plus amino acids, and (3) a minimal medium. Interpret your results if the fungus grows in all three media. Interpret your results if the fungus doesn’t grow in any of the media. 6. EVOLUTION LINK. Because introns are present in all eukaryotes and a few prokaryotes, biologists hypothesize that introns evolved very early in the history of life. If that is true, suggest why the majority of prokaryotes living today lost most of their introns during the course of evolution. Additional questions are available in CengageNOW at www.cengage.com/ login.

14

Gene Regulation

Courtesy of David J. Goldhamer, University of Connecticut

Specific gene expression in a mouse embryo. This 13.5-dayold mouse embryo has been genetically engineered so that cells that transcribe and express the myogenin gene stain blue. The myogenin gene is expressed only in those cells of the embryo that will give rise to muscle tissue. (The gestation period for a mouse is 18 to 20 days.)

KEY CONCEPTS

E

ach type of cell in a multicellular organism has a characteristic shape, carries out very specific activities, and makes a distinct set of proteins.

14.1 Cells regulate which genes will be expressed and

Yet with few exceptions, all cells in an organism contain the same genetic

when. Gene regulation in bacteria is primarily in response to stimuli in the environment; gene regulation in eukaryotes helps to maintain homeostasis.

information. Why, then, aren’t cells identical in structure, molecular com-

14.2 Gene regulation in bacteria occurs primarily at the level of transcription; regulation of transcription can be either positive or negative.

position, and function? Cells differ because gene expression is regulated, and only certain subsets of the total genetic information are expressed in any given cell (see photograph). What mechanisms control gene expression? Consider a gene coding

14.3 Gene regulation in eukaryotes occurs at the lev-

for a protein that is an enzyme. Expressing that gene involves three basic

els of transcription, posttranscription, translation, and posttranslation.

steps: transcribing the gene to form messenger RNA (mRNA), translating the mRNA into protein, and activating the protein so it can catalyze

series of processes, each of which is regulated in many ways. The control mechanisms use various signals, some originating within the cell and others coming from other cells or from the environment. These signals interact with DNA, RNA, or protein. Mechanisms that regulate gene expression include control of the amount of mRNA transcribed, of the rate of translation of mRNA, and of the activity of the protein product. These controls are accomplished in several ways. For example, the rate of transcription and the rate of mRNA degradation both control the amount of available mRNA. Although bacteria are not multicellular, regulation of gene expression is also essential for their survival. Gene regulation in bacteria often involves controlling the transcription of genes whose products are involved in resource use. In eukaryotes, by contrast, fine-tuning the control systems occurs at all levels of gene regulation. Involving all levels of gene regulation is consistent with the greater complexity of eukaryotic cells and the need for developmental controls in multicellular organisms.

14.1 GENE REGULATION IN BACTERIA AND EUKARYOTES: AN OVERVIEW ■ ■ LEARNING OBJECTIVE 1

Explain why bacterial and eukaryotic cells have different mechanisms of gene regulation.

Bacterial and eukaryotic cells use distinctly different mechanisms of gene regulation, based on the specific requirements of the organism. Bacterial cells can exist independently, and each cell performs all its own essential functions. Because they grow rapidly and have relatively short times between cell division events, bacteria carry fewer chemical components. Expression of bacterial genes occurs when it helps bacteria adjust to their environment, including any food that might be present in the surroundings. The primary requirement of bacterial gene regulation is the production of enzymes and other proteins when needed, and transcriptional-level control is the most efficient mechanism. The organization of related genes into groups that are rapidly turned on and off as units allows the synthesis of only the gene products needed at any particular time. This type of regulation requires rapid turnover of mRNA molecules to prevent messages from accumulating and continuing to be translated when they are not needed. Bacteria rarely regulate enzyme levels by degrading proteins. Once the synthesis of a protein ends, the previously synthesized protein molecules are diluted so rapidly in subsequent cell divisions that breaking them down is usually not necessary. Only when cells are starved or deprived of essential amino acids do

protein-digesting enzymes recycle amino acids by breaking down proteins no longer needed for survival. Eukaryotic cells have different regulatory requirements. Although transcriptional-level control predominates, control at other levels of gene expression is also very important, especially in multicellular organisms, in which groups of cells cooperate with one another in a division of labor. Because a single gene is regulated in different ways in different types of cells, eukaryotic gene regulation is complex. Eukaryotic cells usually have a long lifespan during which they may need to maintain homeostasis while responding to many different stimuli. New enzymes are not necessarily synthesized each time the cells respond to a stimulus; instead, in many instances pre-formed enzymes and other proteins are rapidly converted from an inactive to an active state. Some cells have a large store of inactive messenger RNA; for example, the mRNA of an egg cell becomes activated when the egg is fertilized. Much gene regulation in multicellular organisms is focused on the differential expression of genes in the cells in various tissues. Each type of cell has certain genes that are active and other genes that may never be used. For example, developing red blood cells produce the oxygen transport protein hemoglobin, whereas muscle cells never produce hemoglobin but instead produce myoglobin, a related protein coded for by a different gene that stores oxygen in muscle tissues. Apparently, the selective advantages of cell cooperation in multicellular eukaryotes far outweigh the detrimental effects of carrying a load of inactive genes through many cell divisions. The domesticated pig provides another example of how gene regulation in multicellular organisms affects expression in different tissues. In 2003, biologists reported details of a genetic mutation in pigs that makes them develop more muscle tissue (FIG. 14-1). The mutation is in a gene, designated insulin-like growth factor

H. Schwind/Okapia/Science Photo Library/Photo Researchers, Inc.

a specific reaction in the cell. Thus, gene expression results from a

FIGURE 14-1 A lean pig with a mutation in the IGF2 gene Years ago, farmers discovered a pig with more muscle and less fat. They selectively bred this pig, and today most commercial pig populations have the mutation that confers this trait. The mutation, which was recently identified as a substitution of a guanine nucleotide for an adenine nucleotide, occurs in a nonprotein-coding region of the IGF2 gene.

2 (IGF2), which codes for a protein produced by both muscle and liver tissues. The biologists found that a base-substitution mutation (see Chapter 13) in the IGF2 gene makes the gene 3 times more active in pig muscles, resulting in leaner meat. However, the mutation does not change expression of the gene in liver cells. Interestingly, this mutation is not in the protein-coding portion of the gene but in a nonprotein-coding, and presumably regulatory, region.

Review ■ ■

What is the most efficient mechanism of gene regulation in bacteria? How would you contrast gene regulation in prokaryotes with that in eukaryotes?

14.2 GENE REGULATION IN BACTERIA ■ ■ LEARNING OBJECTIVES 2

Define operon and explain the functions of the operator and promoter regions. Distinguish among inducible, repressible, and constitutive genes.

3 4 Differentiate between positive and negative control, and show how both 5

types of control operate in regulating the lac operon. Describe the types of posttranscriptional control in bacteria.

The Escherichia coli bacterium is common in the intestines of humans and other mammals. It has 4288 genes that code for proteins, approximately 60% of which have known functions. Some of these genes encode proteins that are always needed (such as enzymes involved in glycolysis). These genes, which are constantly transcribed, are constitutive genes, and biologists describe them as constitutively expressed. Other proteins are needed only when the bacterium is growing under certain conditions. For instance, bacteria living in the colon of an adult cow are not normally exposed to the milk sugar lactose, a disaccharide. If those cells ended up in the colon of a calf, however, they would have lactose available as a source of energy. This situation presents a dilemma. Should a bacterial cell invest energy and materials to produce lactose-metabolizing enzymes just in case it ends up in the digestive system of a calf? Yet if E. coli cells could not produce those enzymes, they might starve in the middle of an abundant food supply. Thus, E. coli functions by regulating many enzymes to efficiently use available organic molecules. Cell metabolic activity is controlled in two ways: by regulating the activity of certain enzymes (how effectively an enzyme molecule works) and/or by regulating the number of enzyme molecules present in each cell. Some enzymes are regulated in both ways. An E. coli cell growing on glucose needs about 800 different enzymes. Some are present in large numbers, whereas only a few molecules of others are required. For the cell to function properly, the quantity of each enzyme must be efficiently controlled. Bacteria respond to changing environmental conditions. If lactose is added to a culture of E. coli cells, they rapidly synthesize the three enzymes needed to metabolize lactose. Bacteria respond so efficiently because functionally related genes—such as the three

genes involved in lactose metabolism—are regulated together in gene complexes called operons.

Operons in bacteria facilitate the coordinated control of functionally related genes The French researchers François Jacob and Jacques Monod are credited with the first demonstration, in 1961, of gene regulation at the biochemical level through their elegant studies on the genes that code for the enzymes that metabolize lactose. In 1965, they received the Nobel Prize in Physiology or Medicine for their discoveries relating to genetic control of enzymes. To use lactose as an energy source, E. coli cells first cleave the sugar into the monosaccharides glucose and galactose, using the enzyme b-galactosidase. Another enzyme converts galactose to glucose, and enzymes in the glycolysis pathway further break down the resulting two glucose molecules (see Chapter 8). E. coli cells growing on glucose produce very little bgalactosidase enzyme. However, each cell grown on lactose as the sole carbon source has several thousand b-galactosidase molecules, accounting for about 3% of the cell’s total protein. Amounts of two other enzymes, lactose permease and galactoside transacetylase, also increase when the cells are grown on lactose. The cell needs permease to transport lactose efficiently across the bacterial plasma membrane; without it, only small amounts of lactose enter the cell. The transacetylase may function in a minor aspect of lactose metabolism, although its role is not clear. Jacob and Monod identified mutant strains of E. coli in which a single genetic defect wiped out all three enzymes. This finding, along with other information, led the researchers to conclude that the DNA coding sequences for all three enzymes are linked as a unit on the bacterial DNA and are controlled by a common mechanism. Each enzyme-coding sequence is a structural gene. Jacob and Monod coined the term operon for a gene complex consisting of a group of structural genes with related functions plus the closely linked DNA sequences responsible for controlling them. The structural genes of the lac operon (lactose operon)—lacZ, lacY, and lacA—code for b-galactosidase, lactose permease, and transacetylase, respectively (FIG. 14-2). Transcription of the lac operon begins as RNA polymerase binds to a single promoter region upstream from the coding sequences. It then proceeds to transcribe the DNA, forming a single mRNA molecule that contains the coding information for all three enzymes. Each enzyme-coding sequence on this mRNA contains its own start and stop codons; thus, the mRNA is translated to form three separate polypeptides. Because all three enzymes are translated from the same mRNA molecule, their synthesis is coordinated by turning a single molecular “switch” on or off. The switch that controls mRNA synthesis is the operator, which is a sequence of bases upstream from the first structural gene in the operon. In the absence of lactose, a repressor protein called the lactose repressor binds tightly to the operator. RNA polymerase binds to the promoter but is blocked from transcribing the protein-coding genes of the lac operon. The lactose repressor protein is encoded by a repressor gene, which in this case is an adjacent structural gene located upstream

KEY POINT

The structural genes of the lac operon are coordinately controlled and transcribed as a single mRNA. lac operon

Repressor gene

Promoter Operator

DNA Transcription

lacY lacA

Repressor protein

mRNA Ribosome

lacZ

Translation

Precursor of repressor protein

(a) Lactose absent. In the absence of lactose, a repressor protein, encoded by an adjacent gene, binds to a region known as the operator, thereby blocking transcription of the structural genes.

Repressor gene

mRNA

lac operon Promoter Operator

lacZ

lacY lacA

Transcription

RNA polymerase mRNA

Inducer (allolactose) Repressor protein (inactive)

Transacetylase

Translation

Lactose permease β-galactosidase Enzymes for lactose metabolism

(b) Lactose present. When lactose is present, it is converted to allolactose, which binds to the repressor at an allosteric site, altering the structure of the protein so it no longer binds to the operator. As a result, RNA polymerase is able to transcribe the structural genes.

FIGURE 14-2 Animated The lac operon

from the promoter site. Unlike the lac operon genes, the repressor gene is constitutively expressed and is therefore constantly transcribed; the cell continuously produces small amounts of the repressor protein. The repressor protein binds specifically to the lac operator sequence. When cells grow in the absence of lactose, a repressor molecule nearly always occupies the operator site. When the operator site is briefly free of the repressor, the cell synthesizes a small amount of mRNA. However, the cell synthesizes very few enzyme molecules because E. coli mRNA is degraded rapidly (it has a halflife of about 2 to 4 minutes). Lactose “turns on,” or induces, the transcription of the lac operon because the lactose repressor protein contains a second functional region (an allosteric site; see Chapter 7) separate from its DNA-binding site. This allosteric site binds to allolactose, a struc-

tural isomer made from lactose. If lactose is in the growth medium, a few molecules enter the cell and are converted to allolactose by the few b-galactosidase molecules present. The binding of a molecule of allolactose to the repressor alters the shape of the protein so that its DNA-binding site no longer recognizes the operator. When the repressor molecules have allolactose bound to them and are therefore inactivated, RNA polymerase actively transcribes the structural genes of the operon. The E. coli cell continues to produce b-galactosidase and the other lac operon enzymes until it uses up virtually all the lactose. When the intracellular level of lactose drops, so too does the concentration of allolactose. Therefore, allolactose dissociates from the repressor protein, which then assumes a shape that lets the repressor bind to the operator region and shut down transcription of the operon. Jacob and Monod isolated genetic mutants to study the lac operon How did Jacob and Monod elucidate the functioning of the lac operon? Their approach involved the use of mutant strains, which even today are an essential tool of researchers trying to unravel the components of various regulatory systems. Genetic crosses of mutant strains allow investigators to determine the map positions (linear order) of the genes on the DNA and to infer normal gene functions by studying what happens when they are missing or altered. Researchers usually combine this information with the results of direct biochemical studies. Jacob and Monod divided their mutant strains into two groups, based on whether a particular mutation affected only one enzyme or all three (FIG. 14-3). In one group, only one enzyme of the three—b-galactosidase, lactose permease, or galactoside transacetylase—was affected. Subsequent gene-mapping studies showed these were mutations in structural genes located next to each other in a linear sequence. Jacob and Monod also studied regulatory mutants in which a single mutation affected the expression of all three enzymes. Some of the regulatory mutants transcribed the structural genes of the lac operon at a significant rate, even in the absence of lactose, causing the cell to waste energy in producing unneeded enzymes. One group of these gene mutations had map positions just outside the lac operon. Jacob and Monod hypothesized the existence of a repressor gene that codes for a repressor molecule (later found to be a protein). Although the specific defect may vary, the members of this group of mutants produce defective repressors; hence, no binding to the lac operator and promoter takes place, and the lac operon is constitutively expressed. In contrast, some regulatory mutants failed to transcribe the lac operon even when lactose was present. Researchers eventually found that each of these abnormal genes coded for a repressor with an abnormal binding site that prevented allolactose from binding, although the repressor could still bind to the operator. Once bound to the operator, such a mutant repressor remained bound, keeping the operon “turned off.” The genes responsible for another group of regulatory mutants had map positions within the lac operon but did not directly involve any of the three structural genes. Jacob and Monod hypothesized that the members of this group produced normal repressor mol-

KEY POINT

Jacob and Monod analyzed the properties of various mutant strains of E. coli to deduce the structure and function of the lac operon. Mutation affects one or all three enzymes involved in lactose metabolism Only one enzyme affected

All three enzymes affected equally CONCLUSION: Mutation affects gene regulation

CONCLUSION: Mutation is in a structural gene

Enzymes not present, even when lactose is present

Enzymes always present, even when lactose is absent

CONCLUSION: Mutant strain is not constitutive

CONCLUSION: Mutant strain is constitutive

β-galactosidase absent

Lactose permease absent

Galactoside transacetylase absent

Mutation located outside lac operon

Mutation located outside lac operon

Mutation located within lac operon

CONCLUSION: Mutation in lacZ

CONCLUSION: Mutation in lacY

CONCLUSION: Mutation in lacA

CONCLUSION: Defective repressor cannot bind to inducer but can bind to operator

CONCLUSION: Defective repressor cannot bind to operator

CONCLUSION: Normal repressor cannot bind to defective operator

FIGURE 14-3 Genetic and biochemical characterization of the lac operon The diagram shows how Jacob and Monod inferred conclusions based on their experimental results.

ecules but had abnormal operator sequences incapable of binding to the repressor. An inducible gene is not transcribed unless a specific inducer inactivates its repressor

Geneticists call the lac operon an inducible operon. A repressor usually controls an inducible gene or operon by keeping it “turned off.” The presence of an inducer (in this case, allolactose) inactivates the repressor, permitting the gene or operon to be transcribed. Inducible genes or operons usually code for enzymes that are part of catabolic pathways, which break down molecules to provide both energy and components for anabolic reactions. This type of regulatory system lets the cell save the energy cost of making enzymes when no substrates are available on which they can act. A repressible gene is transcribed unless a specific repressor–corepressor complex is bound to the DNA Another type of gene regulation system in bacteria is associated mainly with anabolic pathways such as those in which cells synthesize amino acids, nucleotides, and other essential biological molecules from simpler materials. Enzymes coded by repressible genes normally regulate these pathways. Repressible operons and genes are usually “turned on”; they are turned off only under certain conditions. In most cases the mo-

lecular signal for regulating these genes is the end product of the anabolic pathway. When the supply of the end product (such as an amino acid) becomes low, all enzymes in the pathway are actively synthesized. When intracellular levels of the end product rise, enzyme synthesis is repressed. Because the growing cell continuously needs compounds such as amino acids, the most effective mechanism for the cell is to keep the genes that control their production turned on, except when a large supply of the amino acid is available. The ability to turn the genes off prevents cells from overproducing amino acids and other molecules that are essential but expensive to make in terms of energy. The trp operon (tryptophan operon) is an example of a repressible system. In both E. coli and a related bacterium, Salmonella, the trp operon consists of five structural genes that code for the enzymes that the cell needs to synthesize the amino acid tryptophan; these genes are clustered as a transcriptional unit with a single promoter and a single operator (FIG. 14-4a). A distant repressor gene codes for a diffusible repressor protein, which differs from the lactose repressor in that the cell synthesizes it in an inactive form that cannot bind to the operator region of the trp operon. The DNA-binding site of the repressor becomes effective only when tryptophan, its corepressor, binds to an allosteric site on the repressor (FIG. 14-4b). When intracellular tryptophan levels are low, the repressor protein is inactive and cannot bind to the operator region of the DNA. The enzymes required for tryptophan

KEY POINT

Repressor gene

DNA mRNA

Structural genes coding for enzymes that synthesize the amino acid tryptophan are organized in a repressible operon. trp operon Promoter Operator

RNA polymerase

trpE

trpD

trpC

trpB

trpA

Transcription mRNA Translation

Repressor protein (inactive)

Enzymes of the tryptophan biosynthetic pathway

Tryptophan

(a) Low intracellular tryptophan levels. Repressor protein is unable to prevent transcription because it cannot bind to the operator.

Repressor gene

trp operon Promoter Operator

DNA

trpE

trpD

trpC

trpB

trpA

Active repressor– corepressor complex

mRNA Inactive repressor protein Tryptophan (corepressor)

(b) High intracellular tryptophan levels. The amino acid tryptophan binds to an allosteric site on the repressor protein, changing its conformation. The resulting active form of the repressor binds to the operator region, blocking transcription of the operon until tryptophan is again required by the cell.

FIGURE 14-4 The trp operon

synthesis are produced, and the intracellular concentration of tryptophan increases. Some of it binds to the repressor, altering the repressor’s shape so it binds tightly to the operator. This switches the operon off, thereby blocking transcription. Negative regulators inhibit transcription; positive regulators stimulate transcription The features of the lac and trp operons we have described so far are examples of negative control, a regulatory mechanism in which the DNA binding regulatory protein is a repressor that turns off transcription of the gene. Positive control is regulation by acti-

vator proteins that bind to DNA and thereby stimulate the transcription of a gene. The lac operon is controlled by both a negative regulator (the lactose repressor) and a positively acting activator protein. Positive control of the lac operon requires the cell to recognize the absence of the sugar glucose, which is the initial substrate in the glycolysis pathway. Lactose, like glucose, undergoes stepwise breakdown to yield energy. However, because glucose is a product of the catabolic hydrolysis of lactose, it is most efficient for E. coli cells to use the available supply of glucose first, sparing the cell the considerable energy cost of making additional enzymes such as b-galactosidase (FIG. 14-5a). The lac operon has a very inefficient promoter element—that is, it has a low affinity for RNA polymerase, even when the repressor protein is inactivated. However, a DNA sequence adjacent to the promoter site is a binding site for another regulatory protein, the catabolite activator protein (CAP). When activated, CAP stimulates transcription of the lac operon and several other bacterial operons. In its active form, CAP is bound by an allosteric site to cyclic AMP, or cAMP, an alternative form of adenosine monophosphate. Examine Figure 3-25 to see the chemical structure of cAMP. As the cells become depleted of glucose, cAMP levels increase (FIG. 14-5b). After the cAMP molecules bind to CAP, the resulting active complex then binds to the CAPbinding site near the lac operon promoter. This binding of active CAP bends DNA’s double helix (FIG. 14-6), strengthening the affinity of the promoter region for RNA polymerase so that the rate of transcriptional initiation accelerates in the presence of lactose. Thus, the lac operon is fully active only if lactose is available and intracellular glucose levels are low. TABLE 14-1 summarizes negative and positive controls in bacteria.

Constitutive genes are transcribed at different rates Many gene products encoded by E. coli DNA are needed only under certain environmental or nutritional conditions. As you have seen, these genes are generally regulated at the level of transcription. They are turned on and off as metabolic and environmental conditions change. By contrast, constitutive genes are continuously transcribed, but they are not necessarily transcribed (or their mRNAs translated) at the same rate. Some enzymes work more effectively or are more stable than others and thus are present in smaller amounts. Constitutive genes that encode proteins required in large amounts generally have greater expression—that is, are transcribed more rapidly—than genes coding for proteins re-

KEY POINT

The lactose promoter by itself is weak and binds RNA polymerase inefficiently even when the lactose repressor is inactive. Promoter

Repressor gene

RNA CAP- polymerase– binding binding Operator site site

lacZ

lacY

lacA

DNA mRNA

RNA polymerase binds poorly

quired in smaller amounts. The promoter elements of these genes control their transcriptional rate. Constitutive genes with efficient (“strong”) promoters bind RNA polymerase more frequently and consequently transcribe more mRNA molecules than those with inefficient (“weak”) promoters. Genes coding for repressor or activator proteins that regulate metabolic enzymes are usually constitutive and produce their protein products constantly. Because each cell usually needs relatively few molecules of any specific repressor or activator protein, promoters for those genes tend to be relatively weak.

Some posttranscriptional regulation occurs in bacteria

CAP (inactive)

As you have seen, much of the variability in protein levels in E. coli is determined by transcriptional-level control. However, regulatory mechanisms after transcription, known as posttranscriptional controls, (a) Lactose high, glucose high, cAMP low. When glucose levels are high, cAMP also occur at various levels of gene expression. is low. CAP is in an inactive form and cannot stimulate transcription. Transcription Translational controls are posttranscriptional occurs at a low level or not at all. controls that regulate the rate at which an mRNA Promoter molecule is translated. Because the lifespan of an mRNA molecule in a bacterial cell is very short, one RNA CAP- polymerase– that is translated rapidly produces more proteins than Repressor binding binding one that is translated slowly. Some mRNA molecules Operator lacZ lacY lacA gene site site in E. coli are translated as much as 1000 times faster than other mRNAs. Most of the differences appear DNA to be due to the rate at which ribosomes attach to the RNA polymerase Transcription mRNA and begin translation. binds efficiently mRNA Posttranslational controls act as switches that mRNA activate or inactivate one or more existing enzymes, CAP thereby letting the cell respond to changes in the Galactoside intracellular concentrations of essential molecules, Translation cAMP transacetylase such as amino acids. A common posttranslational Lactose control adjusts the rate of synthesis in a metabolic permease Allolactose β-galactosidase pathway through feedback inhibition. An example Repressor protein (inactive) of feedback inhibition is diagrammed in Figure 7-15. Enzymes for lactose metabolism The end product of a metabolic pathway binds to (b) Lactose high, glucose low, cAMP high. When glucose concentrations are low, an allosteric site on the first enzyme in the pathway, each CAP polypeptide has cAMP bound to its allosteric site. The active form of temporarily inactivating the enzyme. When the first CAP binds to the DNA sequence, and transcription becomes activated. enzyme in the pathway does not function, all the succeeding enzymes are deprived of substrates. Notice FIGURE 14-5 Positive control of the lac operon that feedback inhibition differs from the repression caused by tryptophan discussed previously. In that case, the end product of the pathway (tryptophan) prevented the formation of new enzymes. Feedback inhibition acts as a fine-tuning mechanism that regulates the activity of the existDNA ing enzymes in a metabolic pathway.

cAMP CAP dimer

▲

rtesy of S. C. Schultz, G. C. Shields, and T. A. Steitz, Yale University

Allolactose Repressor protein (inactive)

FIGURE 14-6 The binding of CAP to DNA

This computer-generated image shows the bend formed in the DNA double helix when it binds to CAP. CAP is a dimer consisting of two identical polypeptide chains, each of which binds one molecule of cAMP.

TABLE 14-1

Transcriptional Control in Bacteria*

Negative Control

Result

Inducible genes Repressor protein alone Lactose repressor alone Repressor protein + inducer Lactose repressor + allolactose

Active repressor turns off regulated gene(s) lac operon not transcribed Inactive repressor–inducer complex fails to turn off regulated gene(s) lac operon transcribed

Repressible genes Repressor protein alone Tryptophan repressor alone Repressor protein + corepressor Tryptophan repressor + tryptophan

Inactive repressor fails to turn off regulated gene(s) trp operon transcribed Active repressor–corepressor complex turns off regulated gene(s) trp operon not transcribed

Positive Control

Result

Activator protein alone CAP alone Activator complex CAP + cAMP

Activator alone cannot stimulate transcription of regulated gene(s) Transcription of lac operon not stimulated Functional activator complex stimulates transcription of regulated gene(s) Transcription of lac operon stimulated

*A general description of each type is followed by a specific example.

Review ■

■ ■

■

What is the function of each of the parts of the lac operon (promoter, operator, and structural genes)? What structural features does the trp operon share with the lac operon? Why do scientists define the trp operon as repressible and the lac operon as inducible? How is glucose involved in positive control of the lac operon?

14.3 GENE REGULATION IN EUKARYOTIC CELLS ■ ■ LEARNING OBJECTIVES 6 Discuss the structure of a typical eukaryotic gene and the DNA elements involved in regulating that gene.

7 Give examples of some of the ways eukaryotic DNA-binding proteins bind to DNA.

8 Illustrate how a change in chromosome structure may affect the activity of a gene.

9 Explain how a gene in a multicellular organism may produce different products in different types of cells.

10 Identify some of the types of regulatory controls that operate in eukaryotes after mature mRNA is formed.

Like bacterial cells, eukaryotic cells respond to changes in their environment. In addition, multicellular eukaryotes require regulation modes that let individual cells commit to specialized roles and let groups of cells organize into tissues and organs. These processes are accomplished mainly by regulating transcription, but posttranscriptional, translational, and posttranslational controls are also important. In Chapters 12 and 13, we observed that in

eukaryotes all aspects of information transfer—including replication, transcription, and translation—are more complicated than in prokaryotes. Not surprisingly, this complexity offers more ways to control gene expression. Unlike many bacterial genes, eukaryotic genes are not typically arranged in operon-like clusters. (A notable exception is the nematodes, or roundworms, in which some genes are organized into operons.) However, each eukaryotic gene has specific regulatory sequences that are essential in controlling transcription. Many “housekeeping” enzymes (those needed by virtually all cells) are encoded by constitutive genes, which are expressed in all cells at all times. Some inducible genes have also been found; these respond to environmental threats or stimuli, such as heavy-metal ingestion, viral infection, and heat shock. For example, when a cell is exposed to high temperature, many proteins fail to fold properly. These unfolded proteins elicit a survival response in which heat-shock genes are transcribed and heat-shock proteins are synthesized. Although the functions of most heat-shock proteins in eukaryotic cells are unknown, some are molecular chaperones that help newly synthesized proteins fold into their proper shape (discussed in Chapters 3 and 13). Some genes are inducible only during certain periods in the life of the organism; they are controlled by temporal regulation. Finally, some genes are under tissue-specific regulation. For example, a gene involved in production of a particular enzyme may be regulated by one stimulus (such as a hormone) in muscle tissue, by an entirely different stimulus in pancreatic cells, and by a third stimulus in liver cells. We explore these types of regulation in more detail in Chapters 17 and 49. FIGURE 14-7 summarizes the regulation of gene expression in eukaryotes. You may want to refer to it as you read the following discussion.

Eukaryotic transcription is controlled at many sites and by many regulatory molecules Like bacterial genes, most genes of multicellular eukaryotes are controlled at the transcriptional level. As you will see, various base sequences in the DNA are important in transcriptional control. In addition, regulatory proteins and the way the DNA is organized in the chromosome affect the rate of transcription.

Cytosol Nucleus Regulation of chromatin DNA must be unpacked; heterochromatin cannot be transcribed. Methylated DNA is inaccessible to transcription machinery; demethylated DNA can be transcribed. Regulation of transcription Selective transcription: promoter and enhancer elements in DNA interact with protein transcription factors to activate or repress transcription.

Chromosome organization affects the expression of some genes A chromosome is not only a bearer of genes. Various arrangements of a chromosome’s ordered compoRegulation of mRNA processing nents increase or decrease expression of the genes it Control mechanisms, such as rate of contains. Figure 10-4 shows the organization of a euintron/exon splicing, regulate mRNA processing. karyotic chromosome. In multicellular eukaryotes, only a subset of the genes in a cell is active at any one Alternative splicing of exons produces time. The inactivated genes differ among cell types different proteins from same mRNA. and in many cases are irreversibly dormant. Some of the inactive genes lie in highly compacted chromatin, visible microscopically as densely stained regions of chromosomes during cell division. Nuclear These regions of chromatin remain tightly coiled Regulation of mRNA transport pore and bound to chromosome proteins throughout Controlling access to, or efficiency of, transport through nuclear pores regulates the cell cycle; even during interphase, they are vismRNA transport from nucleus to cytosol. ible as darkly stained fibers called heterochromatin (FIG. 14-8a). Evidence suggests that most of the heterochromatin DNA is not transcribed. For example, Regulation of translation Translational controls determine how often most of the inactive X chromosome of the two X and how long specific mRNA is translated. chromosomes in female mammals is heterochromatic and appears as a Barr body (see Fig. 11-16). Active genes are associated with a more loosely packed chromatin structure called euchromatin Degradation of mRNA Modifications to protein (FIG. 14-8b). The exposure of the DNA in euchromaTranslational controls determine Chemical modifications, such as degree to which mRNA is protected phosphorylation, affect activity of tin lets it interact with transcription factors (discussed from destruction; proteins that bind protein after it is produced. later in the chapter) and other regulatory proteins. to mRNA in cytosol affect stability. Cells may have several ways to change chromatin structure from heterochromatin to euchromatin. Degradation of protein One way is to chemically modify histones, the proSelective degradation targets teins that associate with DNA to form nucleosomes. specific proteins for destruction Each histone molecule has a so-called tail, a string by proteasomes. of amino acids that extends from the DNA-wrapped nucleosome (see Fig. 10-2). Methyl groups, acetyl groups, sugars, and even proteins chemically attach FIGURE 14-7 Gene regulation in eukaryotic cells to the histone tail and may expose or hide genes, turning them on or off. Such chemical modifications of histones are the example, the genes on DNA in nucleosomes where histone tails focus of active research because they appear to influence gene exare flagged with acetyl groups generally tend to be expressed. Bipression and therefore offer promise of new approaches to treat ologists are now trying to determine if specific histone modificacancer and other diseases with a genetic component. Biologists tions are associated with the activity or inactivity of specific genes have found general correlations between gene activity and cerwithin a cell. tain chemical groups attached to or absent from histone tails. For

(a) Inactive region of DNA. Heterochromatin is organized into tightly associated nucleosomes. Heterochromatin: genes silent Chromatin decondensation Nucleosome Histones DNA Transcribed region Euchromatin: genes active

(b) Active region of DNA. Active genes are found in decondensed chromatin called euchromatin. Euchromatin increases the accessibility to RNA polymerase required for transcription. The histones are physically removed from the DNA in the region where transcription occurs.

FIGURE 14-8 Regulation of chromatin: the effect of chromatin structure on transcription

provides a mechanism for epigenetic inheritance. Evidence continues to accumulate that epigenetic inheritance is an important mechanism of gene regulation over the lifetime of an individual. For example, the methylation of genes increases in many people as they age, and decreases in others, perhaps in response to signals from the environment. New phenotypic traits can arise from epigenetic changes despite the fact that the nucleotide sequence of the gene itself has not changed (FIG. 14-9). Epigenetic changes have also been linked to certain human diseases, such as cancer. For example, tumor suppressor genes that inhibit cell division (and cancer) are sometimes inactivated epigenetically, resulting in cancer. Chapter 17 discusses cancer and tumor suppressor genes. Thus, some inherited charac-

© Randy Jirtle/Duke University

Gene inactivation by DNA methylation is an example of epigenetic inheritance Inactive genes of vertebrates and some other organisms typically show a pattern of DNA methylation in which the DNA has been chemically altered by enzymes that add methyl groups to certain cytosine nucleotides in DNA. (The resulting 5-methylcytosine still pairs with guanine in the usual way.) Evidence indicates that certain regulatory proteins selectively bind to methylated DNA and make it inaccessible to RNA polymerase and other proteins involved in transcription. DNA methylation, which is found in inactive regions of the cell’s genome, probably reinforces gene inactivation rather than serves as the initial mechanism to silence genes. Apparently, once a gene has been turned off by some other means, DNA methylation ensures it will remain inactive. For example, the DNA of the inactive X chromosome of a female mammal becomes methylated after the chromosome has become a condensed Barr body (see Chapter 11). When the DNA replicates in a mitotic cell division, each double strand of DNA has one methylated strand and one unmethylated strand. Methylation enzymes add methyls to the new strand, thereby perpetuating the pre-existing methylation pattern. Hence, the DNA continues to be transcriptionally inactive in all descendants of these cells, often for multiple generations. In mammals, DNA methylation maintained in this way accounts for genomic imprinting (also called parental imprinting), in which the phenotypic expression of certain genes is determined by whether a particular allele is inherited from the female or the male parent. Genomic imprinting is associated with some human genetic diseases, such as Prader–Willi syndrome and Angelman syndrome (both discussed in Chapter 16). Genomic imprinting is an example of epigenetics. Epigenetic inheritance involves changes in how a gene is expressed. In contrast, genetic inheritance involves changes in a gene’s nucleotide sequence. Because DNA methylation patterns tend to be passed to successive cell generations during cell division, methylation

FIGURE 14-9 Epigenetic variation in the agouti gene in mice The DNA of these mice is almost identical, yet their coat colors and body shapes vary significantly. These mice differ in the presence or absence of epigenetic “marks,” such as methyl groups on DNA in the region of a gene known as agouti.

Drosophila chorion gene

of the insect body, however, the gene exists as a single copy in the chromosome.

Gene amplification by repeated DNA replication of chorion gene region

Chorion gene in ovarian cell

FIGURE 14-10 Regulation of transcription: gene amplification In Drosophila, multiple replications of a small region of the chromosome amplify the chorion (eggshell) protein genes. Replication is initiated at a discrete chromosome origin of replication (top pink box) for each copy of the gene that is produced. Replication is randomly terminated, resulting in a series of forked structures in the chromosome.

teristics do not depend exclusively on the sequence of nucleotides in DNA. Gene amplification increases the number of copies of a gene A single gene cannot always provide enough copies of its mRNA to meet the cell’s needs. The requirement for high levels of certain products is met if multiple copies of the genes that encode them are present in the chromosome. Genes of this type, whose products are essential for all cells, may occur as multiple copies arranged one after another along the chromosome, in tandemly repeated gene sequences. Histone genes are usually present as multiple copies of 50 to 500 genes in cells of multicellular organisms. Similarly, multiple copies (150 to 450) of genes for rRNA and tRNA are present in cells. Other genes, which are required only by a small group of cells, are selectively replicated in those cells in a process called gene amplification. For example, the Drosophila chorion (eggshell) gene product is a protein made specifically in cells of the female insect’s reproductive tract. These cells make massive amounts of the protein that envelops and protects the zygote. Amplifying the chorion protein gene by DNA replication meets the demand for chorion mRNA. In other words, the DNA in that small region of the chromosome is copied many times (FIG. 14-10). In other cells

DNA double helix

Eukaryotic promoters vary in efficiency, depending on the presence of functional promoter elements

In eukaryotic as well as bacterial cells, the transcription of any gene requires a base pair where transcription begins, known as the transcription initiation site, or start site, plus a sequence of bases, the promoter, to which RNA polymerase binds. The promoter is a relatively short DNA sequence located adjacent to the gene it regulates. In almost all multicellular eukaryotes, the promoter contains an element called the TATA box. The TATA box, which consists of a sequence of DNA nucleotides such as TATAAA is located about 25 to 30 base pairs upstream from the transcription initiation site (FIG. 14-11). Studies show that mutations in the TATA box reduce the rate of transcription. Some eukaryotic genes have promoter elements in addition to the TATA box. Like the TATA box, these promoter elements have a regulatory function and facilitate the expression of the gene. Many eukaryotic promoters have a promoter element of one or more sequences of nucleotide bases located within a short distance (for example, 80 base pairs) upstream of the RNA polymerase–binding site. Mutations in these elements, like mutations in the TATA box, lower the basic level of transcription. Enhancers increase the rate of transcription, whereas silencers decrease the rate of transcription Regulated eukaryotic genes commonly have DNA sequences called enhancers that help form an active transcription initiation complex. In this way, enhancers increase the rate of RNA synthesis, often by several orders of magnitude. Enhancers are remarkable in many ways. Although they are present in all cells, a particular enhancer is functional only in certain types of cells. An enhancer regulates a gene on the same DNA molecule from very long distances—up to thousands of base pairs away from the promoter—and is either upstream or downstream of the promoter it regulates. Furthermore, if an enhancer element is experimentally cut out of the DNA and inverted, it still regulates the gene it normally controls. Many eukaryotic genes are associated with silencers, DNA sequences that can decrease transcription. Like enhancers, silencers are often located long distances from the genes they regulate. As you will see, enhancers and silencers work by interacting with DNA-binding proteins that regulate transcription. TATA box

Upstream

Silencer

Enhancer

Promoter

FIGURE 14-11 Regulation of transcription: eukaryotic elements Eukaryotic regulatory elements include the promoter and various enhancers and silencers. The promoter elements help determine the basic level of transcription that normally occurs, whereas the enhancers (and silencers) have the potential to greatly increase (or decrease) that rate of transcription. (For simplicity, the DNA double helix is represented as a rod.)

Downstream

Gene

Transcription factors are regulatory proteins with several functional domains We previously discussed some DNA-binding proteins that regulate transcription in bacteria. These included the lactose repressor, the tryptophan repressor, and the catabolite activator protein (CAP). Researchers have identified many more DNA-binding proteins that regulate transcription in eukaryotes than in bacteria; these eukaryotic proteins are collectively known as transcription factors. Researchers have identified more than 2000 transcription factors in humans. It is useful to compare regulatory proteins in bacteria and eukaryotes. Many regulatory proteins are modular molecules; that is, they have more than one domain, a region with its own tertiary structure and function. Each eukaryotic transcription factor, like the regulatory proteins of bacteria, has a DNA-binding domain plus at least one other domain that is either an activator or a repressor of transcription for a given gene. Many transcription factors in eukaryotes (and regulatory proteins in bacteria) have similar DNA-binding domains. One example is the helix-turn-helix arrangement, consisting of two a-helical segments. One of these, the recognition helix, inserts into the major groove of the DNA without unwinding the double helix. The other helps hold the first one in place (FIG. 14-12a). The “turn” in the helix-turn-helix is a sequence of amino acids that forms a sharp bend in the molecule. Other transcription factors have DNA-binding domains with multiple “zinc fingers,” loops of amino acids held together by zinc ions. Each loop includes an a-helix that fits into a groove of the DNA (FIG. 14-12b). Certain amino acid functional groups ex-

posed in each finger have been shown to recognize specific DNA sequences. Many transcription factors have DNA-binding domains that function only as pairs, or dimers. Many of these transcription factors are known as leucine zipper proteins because they are held together by the side chains of leucine and other hydrophobic amino acids (FIG. 14-12c). In some cases the two polypeptides that make up the dimer are identical and form a homodimer. In other instances they are different, and the resulting heterodimer may have very different regulatory properties from those of a homodimer. For a simple and speculative example, let us assume that three regulatory proteins—A, B, and C—are involved in controlling a particular set of genes. These three proteins might associate as dimers in six different ways: three kinds of homodimers (AA, BB, and CC) and three kinds of heterodimers (AB, AC, and BC). Such multiple combinations of regulatory proteins greatly increase the number of possible ways that transcription is controlled. Transcription in eukaryotes requires multiple regulatory proteins that are bound to different parts of the promoter. The general transcriptional machinery is a protein complex that binds to the TATA box of the promoter near the transcription initiation site. That complex is required for RNA polymerase to bind, thereby initiating transcription. Both enhancers and silencers become functional when specific transcription factors bind to them. FIGURE 14-13 shows interactions involving an enhancer and a transcription factor that acts as an activator. Each activator has at least two functional domains: a DNA recognition site that usually binds to an enhancer and a gene activation site that contacts the target in the general transcriptional machinery. The DNA between the enhancer and promoter

–

COO Finger 2

Finger 3

Turn

α-helix

Zinc ion

Finger 1

Leucine zipper region

+

NH3

DNA

(a) Helix-turn-helix. This portion of a transcription factor (purple) contains the helix-turn-helix arrangement. The recognition helix is inserted into the groove of the DNA and is connected to a second helix that helps hold it in place by a sequence of amino acids that form a sharp bend.

DNA

(b) Zinc fingers. Regions of certain transcription factors form projections known as “zinc fingers,” which insert into the grooves of the DNA and bind to specific base sequences.

FIGURE 14-12 Regulation of transcription: regulatory proteins In these illustrations, cylinders represent a-helical regions of regulatory proteins and ribbons represent b-pleated sheets.

DNA

(c) Leucine zipper. This leucine zipper protein is a dimer, held together by hydrophobic interactions involving side chains of leucine and other amino acids.

Enhancer

DNA

Target proteins

TATA box

RNA polymerase

(a) Low rate of transcription. This gene is transcribed at a basic level when the general transcriptional machinery, including RNA polymerase, is bound to the promoter. Enhancer

DNA

TATA box

Activator (transcription factor)

(b) High rate of transcription. A transcription factor that functions as an activator binds to an enhancer. The intervening DNA forms a loop, allowing the transcription factor to contact one or more target proteins in the general transcriptional machinery, thereby increasing the rate of transcription. This diagram is highly simplified, and many more target proteins than the two shown are involved.

FIGURE 14-13 Regulation of transcription: the stimulation of transcription by an enhancer

elements forms a loop that lets an activator bound to an enhancer come in contact with the target proteins associated with the general transcriptional machinery. When this contact occurs, the rate of transcription increases.

The mRNAs of eukaryotes have many types of posttranscriptional control

which commonly occurs in the nucleus, can also take place in the cytosol. When the short poly-A tail of an mRNA is elongated, the mRNA becomes activated and is then translated. Some pre-mRNAs are processed in more than one way

Investigators have discovered several forms of regulation involving mRNA processing. In some instances, the same gene produces one type of protein in one tissue and a related but somewhat different type of protein in another tissue. This is possible because many genes produce pre-mRNA molecules that have alternative splicing patterns; that is, they are spliced in different ways depending on the tissue. Typically, such a gene includes at least one segment that can be either an intron or an exon (see Chapter 13). As an intron, the sequence is removed, but as an exon, it is retained. Through alternative splicing, the cells in each tissue produce their own version of mRNA corresponding to the particular gene (FIG. 14-14). For example, this mechanism produces different forms of troponin, a protein that regulates muscle contraction, in different muscle tissues. The stability of mRNA molecules varies

Controlling the lifespan of a particular kind of mRNA molecule permits control over the number of protein molecules translated from it. In some cases, mRNA stability is under hormonal control. This is true for mRNA that codes for vitellogenin, a protein synthesized in the liver of certain female animals, such as frogs and chickens. Vitellogenin is transported to the oviduct, where it is used in forming egg yolk proteins. The hormone estradiol regulates vitellogenin synthesis. When estradiol levels are high, the half-life of vitellogenin mRNA in frog liver is about 500 hours. When researchers deprive cells of estradiol, the half-life of the mRNA drops rapidly, to less than 165 hours. This quickly lowers cell vitellogenin mRNA levels and decreases synthesis of the vitellogenin protein. In addition to affecting mRNA stability, the hormone seems to control the rate at which the mRNA is synthesized.

The half-life of bacterial mRNA is usually minutes long; eukaryotic mRNA, even when it turns over rapidly, is far more stable. Bacterial mRNA Potential splice sites is transcribed in a form that is translated immediately. In contrast, eukaryotic Exon Exon Intron Exon or intron mRNA molecules undergo further mod5 3′ ′ ification and processing before they are pre-mRNA used in protein synthesis. The message is capped, polyadenylated, spliced, and then transported from the nucleus to Alternative splicing the cytoplasm to initiate translation (see Fig. 13-12). These events represent poExon Exon Exon Exon Exon tential control points for translation of 5′ 5′ 3′ the message and the production of its encoded protein. The addition of a poly-A tail to euFunctional mRNA Functional mRNA in tissue B in tissue A karyotic mRNA, for example, appears necessary to initiate translation. Researchers have shown that mRNAs with FIGURE 14-14 Regulation of mRNA processing: alternative splicing long poly-A tails are efficiently translated, In some cases, a pre-mRNA molecule is processed in more than one way to yield two or more mature whereas mRNAs with short poly-A tails mRNAs, each of which encodes a related, but different, protein. In this generalized example, the gene contains a segment that can be an exon in tissue A (left) but an intron in tissue B (right). are essentially dormant. Polyadenylation,

3′

Posttranslational chemical modifications may alter the activity of eukaryotic proteins Another way to control gene expression is by regulating the activity of the gene product. Many metabolic pathways in eukaryotes, as in bacteria, contain allosteric enzymes regulated through feedback inhibition. In addition, after they are synthesized, many eukaryotic proteins are extensively modified. In proteolytic processing, proteins are synthesized as inactive precursors, which are converted to an active form by the removal of a portion of the polypeptide chain. For example, proinsulin contains 86 amino acids. Removing 35 amino acids yields the hormone insulin, which consists of two polypeptide chains containing 30 and 21 amino acids, respectively, linked by disulfide bridges. Chemical modification, by adding or removing functional groups, reversibly alters the activity of an enzyme. One common way to modify the activity of an enzyme or other protein is to add or remove phosphate groups. Enzymes that add phosphate groups are called kinases; those that remove them are phosphatases. For example, the cyclin-dependent kinases discussed in Chapter 10 help control the cell cycle by adding phosphate groups to certain key proteins, causing them to become activated or inactivated. Chemical modifications such as protein phosphorylation also let the cell respond rapidly to certain hormones, or to fast-changing environmental or nutritional conditions. Posttranslational control of gene expression can also involve protein degradation. The amino acid found at the amino end of a protein correlates with whether the protein is long-lived or shortlived. Some proteins, for example, have a 50% turnover of synthesis and degradation every 3 days. Other proteins, such as proteins in the lens of the human eye, may persist for the entire life of the individual. Proteins that are selectively targeted for destruction are covalently bonded to ubiquitin, a small polypeptide tag that contains 76 amino acids. Additional molecules of ubiquitin are added, forming short chains attached to the soon-to-be degraded protein (FIG. 14-15). A protein tagged by ubiquitin is targeted for degradation in a proteasome. Proteasomes are large macromolecular structures that recognize the ubiquitin tags. Proteases (proteindegrading enzymes) associated with proteasomes hydrolyze some of the peptide bonds, degrading the protein into short, nonfunctional peptide fragments. As the protein is degraded, the ubiquitin molecules are released intact, which allows them to be used again.

■ ■

Target protein

Ubiquitin

1

Ubiquitin molecules attach to protein targeted for degradation.

2

Protein enters proteasome.

Ubiquitinylated protein

Proteasome

3

Ubiquitins are released and available for reuse. Protein is degraded into peptide fragments. Peptide fragments

FIGURE 14-15 Degradation of protein: ubiquitin and proteasomes

Review ■

■

■

■

How does regulation of eukaryotic genes differ from regulation of bacterial genes? Why must certain genes in eukaryotic cells be present in multiple copies? How does chromosome structure affect the activity of some eukaryotic genes? How does alternative splicing give rise to different proteins?

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

14.1 (page 308) 1 Explain why bacterial and eukaryotic cells have different mechanisms of gene regulation. ■ Bacterial cells grow rapidly and have a relatively short lifespan. Transcriptional-level control is the most common way for the prokaryotic cell to regulate gene expression. ■ Eukaryotic cells usually have a relatively long lifespan, during which they must maintain homeostasis in response to many different stimuli. Although transcriptional-level control predominates, control at other levels of gene expression is also important.

14.2 (page 309) 2 Define operon and explain the functions of the operator and promoter regions. ■ An operon is a gene complex consisting of a group of structural genes with related functions plus the closely linked DNA sequences responsible for controlling them. ■ Each operon has a single promoter region upstream from the protein-coding regions; the promoter is where RNA polymerase first binds to DNA before transcription begins.

The operator serves as the regulatory switch for transcriptionallevel control of the operon. The binding of a repressor protein to the operator sequence prevents transcription; although RNA polymerase binds to the promoter, it is blocked from transcribing the structural genes. When the repressor is not bound to the operator, transcription proceeds. 3 Distinguish among inducible, repressible, and constitutive genes. ■ An inducible operon, such as the lac operon, is normally turned off. The repressor protein is synthesized in an active form that binds to the operator. If lactose is present, it is converted to allolactose, the inducer, which binds to the repressor protein. The altered repressor cannot bind to the operator, and the operon is turned on. ■

Watch the lactose operon in action by clicking on the figure in CengageNOW. A repressible operon, such as the trp operon, is normally turned on. The repressor protein is synthesized in an inactive form that cannot bind to the operator. A metabolite (usually the end product of a metabolic pathway) acts as a corepressor. When corepressor levels are high, a corepressor molecule binds to the repressor, which can now bind to the operator and turn off transcription of the operon. ■ Constitutive genes are neither inducible nor repressible; they are active at all times. The activity of constitutive genes is controlled by how efficiently RNA polymerase binds to their promoter regions. 4 Differentiate between positive and negative control, and show how both types of control operate in regulating the lac operon. ■ Repressible and inducible operons are under negative control. When the repressor protein binds to the operator, transcription of the operon is turned off. ■ Some inducible operons are also under positive control, in which an activator protein binds to the DNA and stimulates transcription of the gene. Catabolite activator protein (CAP) activates the lac operon; CAP binds to the promoter region, stimulating transcription by binding RNA polymerase tightly. To bind, CAP requires cyclic AMP (cAMP), which increases in the cell as glucose decreases. 5 Describe the types of posttranscriptional control in bacteria. ■ Some posttranscriptional controls operate in bacteria. A translational control is a posttranscriptional control that regulates the rate of translation of a particular mRNA. Posttranslational controls include feedback inhibition of key enzymes in some metabolic pathways. ■

14.3 (page 314) 6 Discuss the structure of a typical eukaryotic gene and the DNA elements involved in regulating that gene. Eukaryotic genes are not normally organized into operons. Regulation of eukaryotic genes occurs at the levels of transcription, mRNA processing, translation, and modifications of the protein product. ■ Transcription of a gene requires a transcription initiation site, where transcription begins, plus a promoter to which RNA polymerase binds. In almost all multicellular eukaryotes, the promoter contains an element called the TATA box that has a regulatory function and facilitates expression of the gene. ■

■

Some eukaryotic genes have enhancers and silencers located thousands of bases away from the promoter. These regulatory elements increase or decrease the rate of transcription.

DNA Upstream double helix Silencer

TATA box Enhancer

Promoter

Downstream Gene

7 Give examples of some of the ways eukaryotic DNA-binding proteins bind to DNA. ■ Eukaryotic genes are controlled by DNA-binding proteins called transcription factors. Many are transcriptional activators; others are transcriptional repressors. ■ Each transcription factor has a DNA-binding domain. Some transcription factors have a helix-turn-helix arrangement and insert one of the helices into the DNA. Other transcription factors have loops of amino acids held together by zinc ions; each loop includes an a-helix that fits into the DNA. Some transcription factors are leucine zipper proteins that associate as dimers that insert into the DNA. 8 Illustrate how a change in chromosome structure may affect the activity of a gene. ■ Densely packed regions of chromosomes, called heterochromatin, contain inactive genes. Active genes are associated with a loosely packed chromatin structure called euchromatin. Methyl groups, acetyl groups, sugars, and proteins may chemically attach to the histone tail, a string of amino acids that extends from the DNA-wrapped nucleosome, and may expose or hide genes, turning them on or off. ■ DNA methylation perpetuates gene inactivation. Epigenetic inheritance is an important mechanism of gene regulation that involves changes in how a gene is expressed. Because DNA methylation patterns tend to be repeated in successive cell generations, they provide a mechanism for epigenetic inheritance. ■ Some genes whose products are required in large amounts exist as multiple copies in the chromosome. In the process of gene amplification, some cells selectively amplify genes by DNA replication. 9 Explain how a gene in a multicellular organism may produce different products in different types of cells. ■ As a result of alternative splicing, a single gene produces different forms of a protein in different tissues, depending on how the pre-mRNA is spliced. Typically, such a gene contains a segment that can be either an intron or an exon. As an intron, the sequence is removed, and as an exon, the sequence is retained. 10 Identify some of the types of regulatory controls that operate in eukaryotes after mature mRNA is formed. ■ Certain regulatory mechanisms increase the stability of mRNA, allowing more protein molecules to be synthesized before mRNA degradation. Sometimes mRNA stability is under hormonal control. ■ Posttranslational control of eukaryotic genes occurs by feedback inhibition or by modification of the protein structure. The function of a protein is changed by kinases adding phosphate groups or by phosphatases removing phosphates. ■ Posttranslational control of gene expression also involves protein degradation. Proteins targeted for destruction are covalently bonded to ubiquitin, which tags it for degradation in a proteasome, a large macromolecular structure. Proteases associated with proteasomes degrade the protein into short peptide fragments.

T E S T YO U R U N D E R S TA N D I N G 1. The regulation of most bacterial genes occurs at the level of (a) transcription (b) translation (c) replication (d) posttranslation (e) postreplication

2. The operator of an operon (a) encodes information for the repressor protein (b) is the binding site for the inducer (c) is the binding site for the repressor protein (d) is the binding site for RNA polymerase (e) encodes the information for the CAP

3. A mutation that inactivates the repressor gene of the lac operon results in (a) the continuous transcription of the structural genes (b) no transcription of the structural genes (c) the binding of the repressor to the operator (d) no production of RNA polymerase (e) no difference in the rate of transcription 4. A repressible operon codes for the enzymes of the following pathway.

A

Enzyme 1

¡ B

Enzyme 2

¡ C

Enzyme 3

¡ D

Which component of the pathway is most likely to be the corepressor for that operon? (a) substance A (b) substance B or C (c) substance D (d) enzyme 1 (e) enzyme 3 5. Feedback inhibition is an example of control at the level of (a) transcription (b) translation (c) posttranslation (d) replication (e) all the preceding 6. Which of the following is an example of positive control? (a) transcription occurs when a repressor binds to an inducer (b) transcription cannot occur when a repressor binds to a corepressor (c) transcription is stimulated when an activator protein binds to DNA (d) a and b (e) a and c

each other (c) one DNA region to another DNA region (d) amino acids to zinc atoms (e) RNA polymerase to the operator 8. Inactive genes tend to be found in (a) highly condensed chromatin, known as euchromatin (b) decondensed chromatin, known as euchromatin (c) highly condensed chromatin, known as heterochromatin (d) decondensed chromatin, known as heterochromatin (e) chromatin that is not organized as nucleosomes 9. Which of the following is characteristic of genes and gene regulation in bacteria, but not in eukaryotes? (a) presence of enhancers (b) capping of mRNAs (c) many chromosomes per cell (d) binding of DNA to regulatory proteins (e) no requirement for exon splicing 10. Which of the following is characteristic of genes and gene regulation in both bacteria and eukaryotes? (a) promoters (b) noncoding DNA within coding sequences (c) enhancers (d) operons (e) DNA located in a nucleus 11. Through alternative splicing, eukaryotes (a) reinforce gene inactivation (b) prevent transcription of heterochromatin (c) produce related but different proteins in different tissues (d) amplify genes to meet the requirement of high levels of a gene product (e) bind transcription factors to enhancers to activate transcription

7. The “zipper” of a leucine zipper protein attaches (a) specific amino acids to specific DNA base pairs (b) two polypeptide chains to

CRITICAL THINKING 1. Compare the types of bacterial genes associated with inducible operons, those associated with repressible operons, and those that are constitutive. Predict the category into which each of the following would most likely fit: (a) a gene that codes for RNA polymerase, (b) a gene that codes for an enzyme required to break down maltose, (c) a gene that codes for an enzyme used in the synthesis of adenine.

Mutant b: The map position of this mutation is in the trp operon. The mutant cells do not produce any enzymes coded for by the trp operon under any conditions.

2. The regulatory gene that codes for the tryptophan repressor is not tightly linked to the trp operon. Would it be advantageous if it were? Explain your answer.

6. ANALYZING DATA. You are studying the rate of transcription of a particular eukaryotic gene. When the DNA located several thousand bases upstream of the gene is removed, the transcription rate of the gene decreases dramatically. How would you interpret these results?

3. EVOLUTION LINK. Suggest why evolution resulted in multiple levels of gene regulation in multicellular eukaryotes. 4. EVOLUTION LINK. Comparing DNA sequences in different species indicates that more DNA segments that do not code for protein have been conserved (unchanged) than protein-coding regions. These nonprotein-coding regions are interpreted as gene regulatory elements. Suggest why gene regulatory elements have not undergone many changes during the course of evolution. 5. ANALYZING DATA. Develop a simple hypothesis that would explain the behavior of each of the following types of mutants in E. coli: Mutant a: The map position of this mutation is in the trp operon. The mutant cells are constitutive; that is, they produce all the enzymes coded for by the trp operon, even if large amounts of tryptophan are present in the growth medium.

Mutant c: The map position of this mutation is some distance from the trp operon. The mutant cells are constitutive; that is, they produce all the enzymes coded for by the trp operon, even if the growth medium contains large amounts of tryptophan.

Promoter

Gene

Additional questions are available in CengageNOW at www.cengage.com/ login.

15

DNA Technology and Genomics

Courtesy of GloFish®

Transgenic fish (GloFish®). These aquarium fish glow green, red, or yellow because researchers inserted a foreign gene encoding a fluorescent protein from either jellyfish or sea corals.

KEY CONCEPTS

B

eginning in the mid-1970s, the development of new ways to study DNA led to radically new research approaches, such as recombinant

15.1 Scientists use DNA technology to produce many cop-

DNA technology, in which researchers splice together DNA from differ-

ies of specific genes (gene cloning).

ent organisms in the laboratory. One goal of this technology is to enable

15.2 Biologists study DNA using gel electrophoresis, DNA blotting, automated sequencing, and other methods.

scientists to obtain many copies of a specific DNA segment for the pur-

15.3 Genomics is an emerging field that comprises the

emerging, we do not attempt to explore them all here. Instead, we discuss

structure, function, and evolution of genomes.

15.4 DNA technology and genomics have wide applications, from medical to forensic to agricultural.

15.5 Scientists must assess the risks of each new recombinant organism.

pose of studying it. Because new methods to analyze DNA are continually some of the major approaches that have provided a foundation for the technologies commonly used by molecular geneticists. We then consider how studies of DNA sequences have helped scientists understand the organization of genes and the relationship between genes and their products. In fact, most of our current knowledge of the structure and control of eukaryotic genes, and of the roles of genes in development, comes from applying these methods. DNA sequencing has also revolutionized systematics by helping to clarify evolutionary relationships. This chapter also explores some of the practical applications of DNA technologies. One of the rapidly advancing areas of study is molecular modification—altering the DNA of an organism to produce new genes with new traits. Molecular modification, also called genetic engineering, can take many forms, ranging from basic research to the production of

strains of bacteria that manufacture useful protein products, to

Restriction enzymes are “molecular scissors”

the development of plants and animals that express foreign genes

Discovering restriction enzymes was a major breakthrough in developing recombinant DNA technology. Today, large numbers of different types of restriction enzymes, each with its own characteristics, are readily available to researchers. For example, a restriction enzyme known as HindIII searches a DNA molecule for the restriction site 5¿¬AAGCTT¬3¿, and when it finds that site, it cuts the DNA. (A restriction site is a DNA sequence containing the cleavage site that is cut by a particular restriction enzyme.) The sequence 5¿¬GAATTC¬3¿ is cut by another restriction enzyme, known as EcoRI. The names of restriction enzymes are generally derived from the names of the bacteria from which they were originally isolated. Thus, HindIII and EcoRI are derived from Hemophilus influenzae and E. coli, respectively. Why do bacteria produce such enzymes? During infection, a bacteriophage injects its DNA into a bacterial cell. The bacterium can defend itself if it has restriction enzymes that can attack the bacteriophage DNA. The bacterial cell protects its own DNA from breakdown by modifying it after replication. An enzyme adds a methyl group to one or more bases in each restriction site so that the restriction enzyme does not recognize and cut the bacterial DNA. Restriction enzymes enable scientists to cut DNA from chromosomes into shorter fragments in a controlled way. Many of the restriction enzymes used for recombinant DNA studies cut palindromic sequences, which means the base sequence of one strand reads the same as its complement when both are read in the 5¿ ¡ 3¿ direction. Thus, in our HindIII example, both strands read 5¿¬AAGCTT¬3¿, which as a double-stranded molecule is diagrammed as follows:

(see photograph). The development of DNA cloning, molecular modification, and related techniques have transformed people’s view of biotechnology, which is the commercial or industrial use of cells or organisms. Today, biotechnology includes numerous applications in such diverse areas as medicine, foods and agriculture, and forensic science.

15.1 DNA CLONING ■ ■ LEARNING OBJECTIVES 1

2

3 4

Explain how a typical restriction enzyme cuts DNA molecules and give examples of the ways in which these enzymes are used in recombinant DNA technology. Distinguish among a genomic DNA library, a chromosome library, and a complementary DNA (cDNA) library; explain why one would clone the same eukaryotic gene from both a genomic DNA library and a cDNA library. Explain how researchers use a DNA probe. Describe how the polymerase chain reaction amplifies DNA in vitro.

Recombinant DNA technology was not developed quickly. It actually had its roots in the 1940s with genetic studies of bacteria and bacteriophages (“bacteria eaters”), the viruses that infect them. (See Figure 12-2, which shows bacteriophages on the bacterium Escherichia coli.) Molecular biologist Paul Berg was the first investigator to construct a recombinant DNA molecule; in 1980, he shared the Nobel Prize in Chemistry for his contributions to recombinant DNA technology. After decades of basic research and the accumulation of extensive knowledge, the technology became feasible and available to the many scientists who now use these methods. In recombinant DNA technology, scientists use restriction enzymes from bacteria to cut DNA molecules only in specific places. Restriction enzymes enable researchers to cut DNA into manageable segments. Each fragment is then incorporated into a suitable vector molecule, a carrier capable of transporting the DNA fragment into a cell. Bacteriophages and DNA molecules called plasmids are two examples of vectors. Bacterial DNA is generally circular; most plasmids are separate, much smaller, circular DNA molecules that may be present and replicate inside a bacterial cell, such as E. coli. Researchers introduce plasmids into bacterial cells by a method called transformation, the uptake of foreign DNA by cells (see Chapter 12). For transformation to be efficient, the researcher alters the cells chemically or by electroporation—delivering an electric shock—to make the plasma membrane permeable to the plasmid DNA molecules. Once a plasmid enters a cell, it is replicated and distributed to daughter cells during cell division. When a recombinant plasmid—one that has foreign DNA spliced into it— replicates in this way, many identical copies of the foreign DNA are made; in other words, the foreign DNA is cloned. Transposons are an efficient vector for transferring genes into mice and other research organisms. Recall from Chapter 13 that a transposon is a mobile genetic element that can cut itself out of one location in a cell’s genome and insert itself into a different location.

5¿¬AAGCTT¬3¿ 3¿¬TTCGAA¬5¿

By cutting both strands of the DNA, but in a staggered fashion, these enzymes produce fragments with identical, complementary, single-stranded ends: 5¿¬A 3¿¬TTCGA

AGCTT¬3¿ A¬5¿

These ends are called sticky ends because they pair by hydrogen bonding with the complementary, single-stranded ends of other DNA molecules that have been cut with the same enzyme (FIG. 15-1). Once the sticky ends of two molecules have been joined in this way, they are treated with DNA ligase, an enzyme that covalently links the two DNA fragments to form a stable recombinant DNA molecule. (DNA ligase was discussed in the section on DNA replication in Chapter 12.)

Recombinant DNA forms when DNA is spliced into a vector In recombinant DNA technology, geneticists cut both the DNA to be cloned and plasmid DNA with the same restriction enzyme. The two DNA samples are then mixed under conditions that facilitate hydrogen bonding between the complementary bases of the sticky ends, and the nicks in the resulting recombinant DNA are sealed by DNA ligase (FIG. 15-2).

5′ 3′

A

A

G

C

T

T

T

T

C

G

A

A

3′ 5′

Plus HindIII restriction enzyme 5′

A T

3′

5′

3′ C

T

G

A

A

G

C

T

5′

3′

T A

Sticky ends

FIGURE 15-1 Animated Cutting DNA with a restriction enzyme Many restriction enzymes, such as HindIII, cut DNA at sequences that are palindromic, producing complementary sticky ends. The small black arrows designate the enzyme’s cleavage sites.

RESEARCH METHOD Why Is It Used?

A plasmid carries foreign DNA into a host bacterial cell, where many copies of the gene of interest are made as the bacterial cell divides. These copies are used in many types of research, for example, to make the gene product encoded by the foreign DNA.

How Is It Done? DNA of interest from another organism

Plasmid from a bacterium

1

Plasmid and DNA from another organism are cut by the same restriction enzyme (in this example, HindIII). This produces molecules with complementary single-stranded ends. Clonable DNA fragment

2

Mix two types of molecules so their sticky ends pair. DNA ligase then forms covalent bonds at junctions, linking fragments.

Recombinant DNA

3

Transfer recombinant DNA molecule to host cell, where it is copied and transcribed to produce gene product.

FIGURE 15-2 Splicing foreign DNA (gene of interest) into a cloning vector (in this example, a bacterial plasmid) to make recombinant DNA

3′ 5′

Bacterial chromosome

AatI Bacterium XbaI

Plasmid

E. coli origin of replication

tr es acyc ist line anc e

Te r

PvuII

SalI

BamHI

-3

re Am s

Y of ea re

g in or i n st licatio p

illin pic ance t is

UR

A

HpaI

ClaI

SmaI

(a) This plasmid vector has many useful features. Researchers constructed it from DNA fragments they had isolated from plasmids, E. coli genes, and yeast genes. The two origins of replication, one for E. coli and one for yeast, Saccharomyces cerevisiae, let it replicate independently in either type of cell. Letters on the outer circle designate sites for restriction enzymes that cut the plasmid only at that position. Resistance genes for the antibiotics ampicillin and tetracycline and the yeast URA-3 gene are also shown. The URA-3 gene is useful when transforming yeast cells lacking an enzyme required for uracil synthesis. Cells that take up the plasmid grow on a uracil-deficient medium.

FIGURE 15-3 Plasmids

The plasmids now used in recombinant DNA work have been extensively manipulated in the laboratory to include features helpful in isolating and analyzing cloned DNA (FIG. 15-3). Among these are an origin of replication (see Chapter 12), one or more restriction sites, and genes that let researchers select cells transformed by recombinant plasmids. These genes allow transformed cells to grow under specified conditions that do not allow untransformed cells to grow. In this way, the researchers use features that are also common in naturally occurring plasmids. Typically, plasmids do not contain genes essential to the bacterial cells under normal conditions. However, they often carry genes that are useful under specific environmental conditions, such as genes that confer resistance to particular antibiotics or that let the cells use a specific nutrient. For example, cells transformed with a plasmid that includes a gene for resistance to the antibiotic tetracycline can grow in a medium that contains tetracycline, whereas untransformed cells cannot. A limiting property of any vector is the size of the DNA fragment it can effectively carry. The size of a DNA segment is often given in kilobases (kb), with 1 kb being equal to 1000 base pairs. Fragments that are smaller than 10 kb are usually inserted into plasmids for use in E. coli. However, larger fragments require the use of bacteriophage vectors, which can handle up to 23 kb of DNA. Other vectors are cosmid cloning vectors, which are combination vectors with features from both bacteriophages and plasmids, and bacterial artificial chromosomes (BACs), which accommodate much larger fragments of DNA. A single BAC can include up

(b) The relative sizes of a plasmid and the main DNA of a bacterium.

to about 200 kb of extra DNA, a feature that made BACs especially useful in the Human Genome Project, which is discussed later in the chapter. Recombinant DNA can also be introduced into cells of eukaryotic organisms. For example, geneticists use modified viruses as vectors in mammal cells. These viruses are disabled so they do not kill the cells they infect. Instead, the viral DNA molecules, as well as any foreign DNA they carry, become incorporated into the cell’s chromosomes after infection. As discussed later, other methods do not require a biological vector.

DNA can be cloned inside cells Because a single gene is only a small part of the total DNA in an organism, isolating the piece of DNA containing that particular gene is like finding a needle in a haystack: a powerful detector is needed. Today, many methods enable biologists to isolate a specific nucleotide sequence from an organism. We start by discussing methods in which DNA is cloned inside bacterial cells. We use the cloning of human DNA as an example, although the procedure is applicable for any organism. A genomic DNA library contains fragments of all DNA in the genome The total DNA in a cell is called its genome. For example, if DNA is extracted from human cells, we refer to it as human genomic DNA. A genomic DNA library is a collection of thousands of DNA fragments that represent all of the DNA in the genome. Each fragment is inserted into a plasmid, which is usually incorporated into a bacterial cell. Thus, a human genomic DNA library is stored in a collection of recombinant bacteria, each with a different fragment of human DNA. Scientists use genomic DNA libraries to isolate and study specific genes. In addition, genomic DNA libraries contain a huge number of sequences that do not code for protein. Individual chromosomes can also be isolated to make a chromosome library containing all the DNA fragments in that specific chromosome. If a gene of interest is known to be associated with a particular chromosome, it is easier to isolate that gene from a chromosome library than from a genomic DNA library. The basic cloning methods just described are used to make genomic DNA or chromosome libraries. First, the DNA is cut with a restriction enzyme, generating a population of DNA fragments (FIG. 15-4). These fragments vary in size and in the genetic information they carry, but they all have identical sticky ends. Ge-

RESEARCH METHOD Why Is It Used?

A genomic library is a collection of all the DNA fragments in an organism’s genome; a chromosome library is a collection of all the DNA fragments in a specific chromosome. Each DNA fragment is “stored” in a clone of recombinant bacterial cells. Scientists use genomic and chromosome libraries to isolate and study genes. Sites of cleavage

How Is It Done? Fragment 1

Fragment 2

Human DNA

Fragment 3

1

Fragment 4

Human DNA is cut with a restriction enzyme, generating a population of DNA fragments.

Produce recombinant DNA

Gene for resistance to antibiotic

R

R

R

2

DNA ligase combines human DNA fragments and bacterial plasmids that already contain a gene for antibiotic resistance.

3

Recombinant plasmids are inserted into antibiotic-sensitive bacteria.

R

Transformation

Bacterium without plasmid 4

Bacteria without plasmid fail to grow.

Bacteria with plasmid live and multiply to form a colony.

Bacteria are incubated on a nutrient medium that also contains antibiotic. Only those cells grow that have incorporated a recombinant plasmid.

Plate with antibiotic-containing medium

FIGURE 15-4 Producing a genomic DNA library or a chromosome library

neticists treat plasmid DNA that will be used as a vector with the same restriction enzyme, which converts the circular plasmids into linear molecules with sticky ends complementary to those of the human DNA fragments. Recombinant plasmids are produced by mixing the two kinds of DNA (human and plasmid) under conditions that promote hydrogen bonding of complementary bases. Then, DNA ligase is used to covalently bond the paired ends of the plasmid and human DNA, forming recombinant DNA. Unavoidably, nonrecombinant plasmids also form (not shown in figure) because some plasmids revert to their original circular shape without incorporating foreign DNA. The geneticists insert plasmids into antibiotic-sensitive bacterial cells by transformation. Because the ratio of plasmids to cells is kept very low, it is rare for a cell to receive more than one plasmid molecule, and not all cells receive a plasmid. The researchers incubate normally antibiotic-sensitive cells on a nutrient medium that includes

antibiotics; the only cells that are able to grow have incorporated a plasmid that contains a gene for antibiotic resistance. In addition, the plasmid has usually been modified in ways that enable researchers to identify those cells containing recombinant plasmids. Genomic DNA and chromosome libraries contain redundancies; that is, certain human DNA sequences have been inserted into plasmids more than once, purely by chance. However, each individual recombinant plasmid—analogous to a book in the library— contains only a single fragment of the total human genome. To identify a plasmid containing a sequence of interest, the researcher clones each plasmid until there are millions of copies to work with. This process occurs as the bacterial cells grow and divide. A dilute sample of the bacterial culture is spread on solid growth medium (agar plates), so the cells are widely separated. Each cell divides many times, yielding a visible colony, which is a clone of genetically identical cells originating from a single cell.

sequence of interest. A common approach to detecting a specific DNA sequence involves using a DNA probe, which is a segment of DNA that is homologous to part of the sequence of interest. The DNA probe is usually a radioactively labeled segment of singlestranded DNA that can hybridize—attach by base pairing—to complementary base sequences in the target DNA. To find a specific DNA, biologists transfer the genomic library to a nitrocellulose or nylon membrane (FIG. 15-5). They treat the cells on the membrane chemically to lyse them, releasing the DNA and making the DNA single-stranded. (Single-stranded DNA

All the cells of a particular colony contain the same recombinant plasmid, so during this process a specific sequence of human DNA is also cloned. The major task is to determine which colony out of thousands contains a cloned fragment of interest. Specific DNA sequences are identified in various ways. A complementary DNA probe detects a specific DNA sequence Suppose a researcher wishes to screen the thousands of recombinant DNA molecules in bacterial cells to find the specific DNA RESEARCH METHOD Why Is It Used?

A radioactive nucleic acid probe reveals the presence of complementary sequences of DNA in a genomic library.

How Is It Done?

1

All clones of the genomic library are stored in multiwell plates. An organism’s entire genomic library is stored in many of these plates, which are stacked with a lid on top.

2

Cells from each clone are transferred to a specific location on a nylon or nitrocellulose membrane. The membrane is treated to break open the bacterial cells, releasing plasmids containing DNA fragments.

3

Labeled probe is added to membrane and hybridizes with any DNA with complementary nucleotide sequences (in this case, the gene of interest.)

4

The hybridization is visualized as a dark spot by placing the membrane against X-ray film, which is then developed.

1 2 A B C

Membrane

Radioactively labeled, single-stranded DNA probe for gene of interest

Membrane

Bag Location of plasmid with gene of interest

FIGURE 15-5 Identifying a specific DNA sequence in a genomic library by hybridization with a nucleic acid probe

binds to the membrane.) Then they incubate the membrane with the radioactive probe mixture to let the probes hybridize with any complementary strands of DNA that may be present. Each spot on the membrane containing DNA that is complementary to that particular probe becomes radioactive and is detected by autoradiography (see Chapter 2), using X-ray film. Each spot on the film therefore identifies a colony containing a plasmid that includes the DNA of interest. The biologist then returns to the genomic library and removes cells that contain the cloned sequence of interest. These cells are then grown in culture to produce millions of new bacteria, all containing the recombinant DNA of interest.

A cDNA library is complementary to mRNA and does not contain introns Researchers frequently wish to clone intact genes while avoiding introns, which are regions of eukaryotic genes that do not code for proteins. Scientists also may wish to clone only genes that are expressed in a particular cell type. In such cases, they construct libraries of DNA copies of mature mRNA from which introns have been removed. The copies, known as complementary DNA (cDNA) because they are complementary to mRNA, also lack introns. The researchers use the enzyme reverse transcriptase (see Chapter 13) to synthesize single-stranded cDNA. DNA polymerase is then used to make the cDNA double-stranded (FIG. 15-6). A cDNA library is formed using mRNA from a single cell type as the starting material. The double-stranded cDNA molecules are inserted into plasmid or virus vectors, which then multiply in bacterial cells. Analyzing cDNA clones lets investigators determine certain characteristics of the protein encoded by the gene, including its amino acid sequence. They can also study the structure of the mature mRNA. Because the cDNA copy of the mRNA does not contain intron sequences, comparing the DNA base sequences in cDNA and the DNA in genomic DNA or chromosome libraries reveals the locations of intron and exon coding sequences in the gene. Cloned cDNA sequences are also useful when geneticists want to produce a eukaryotic protein in bacteria. When they introduce an intron-containing human gene, such as the gene for human growth hormone, into a bacterium, it cannot remove the introns from the transcribed RNA to make a functional mRNA for producing its protein product. If they insert a cDNA clone of the gene into the bacterium, however, its transcript contains an uninterrupted coding region. A functional protein is synthesized if the geneticist inserts the gene downstream of an appropriate bacterial promoter and if he or she places the appropriate bacterial translation initiation sequences in it.

1

Formation of cDNA relies on RNA processing that occurs in the nucleus to yield mature mRNA. Exon

Intron

Exon

DNA in a eukaryotic chromosome

Pre-mRNA

Intron

Exon

Transcription

RNA processing (remove introns)

Mature mRNA 2

Mature mRNA is extracted and purified. Reverse transcriptase is added to tube to produce a single-stranded cDNA complementary to the mRNA.

Reverse transcriptase Mature mRNA

mRNA

cDNA copy of mRNA

3

Specific enzymes then degrade the mRNA.

Degraded RNA

cDNA 4

DNA polymerase is used to synthesize a second strand of DNA.

DNA polymerase 5

The end result is a double-stranded cDNA molecule.

Double-stranded cDNA

The polymerase chain reaction amplifies DNA in vitro

FIGURE 15-6 Animated The formation of cDNA

The methods just described for amplifying, or making multiple copies of, a specific DNA sequence involve cloning DNA in cells, usually those of bacteria. These processes are time consuming and require an adequate DNA sample as the starting material. The polymerase chain reaction (PCR), which biochemist Kary Mullis developed in 1985, lets researchers amplify a tiny sample

of DNA without cloning in a cell. The DNA is amplified millions of times in a few hours. In 1993, Mullis received the Nobel Prize in Chemistry for his work. In PCR, DNA polymerase uses nucleotides and primers to replicate a DNA sequence in vitro (outside a living organism),

researchers use a heat-resistant DNA polymerase, known as Taq polymerase. The name of this enzyme reflects its source, Thermus aquaticus, a bacterium that lives in hot springs in Yellowstone National Park. Because the water in this environment is close to the boiling point, all enzymes in T. aquaticus have evolved to be stable at high temperatures. (Recall from Chapter 3 that, when heated, most proteins are irreversibly denatured—they change their overall shape and lose their biological activity.) Bacteria living in deep-sea thermal vents have evolved similar heat-resistant enzymes.

thereby producing two DNA molecules (FIG. 15-7). Then the researchers denature, or separate by heating, the two strands of each molecule and replicate them again, yielding four double-stranded molecules. After the next cycle of heating and replication, there are eight molecules, and so on, with the number of DNA molecules doubling in each cycle. After only 20 heating and cooling cycles are done using automated equipment, this exponential process yields 220, or more than 1 million, copies of the target sequence! Because the reaction can be carried out efficiently only if the DNA polymerase can remain stable through many heating cycles,

RESEARCH METHOD Why Is It Used?

PCR takes an extremely small sample of DNA and amplifies it to millions of copies in a few hours. 3′5′

5′ 3′ 3′

5′ 3′

5′

Primer 3′ Primer

5′

1

DNA is denatured (separated into single strands) by heat.

2

Primers attach to primer-binding site on each DNA strand.

3

Each DNA strand acts as template for DNA synthesis catalyzed by Taq DNA polymerase. The number of DNA molecules doubles each time cycle is repeated.

3′

Primers Cycle II

Primers

5′

Cycle I

Original doublestranded DNA

etc.

How Is It Done?

The initial reaction mixture includes a very small amount of double-stranded DNA, DNA precursors (deoxyribonucleotides), specific nucleic acid primers, and heat-resistant Taq DNA polymerase. PCR involves repeated cycles of denaturation, primer attachment, and strand elongation by enzymes using complementary base pairing.

FIGURE 15-7 Animated Amplification of DNA by the polymerase chain reaction (PCR)

The PCR technique has virtually limitless applications. It makes cloning easier. It enables researchers to amplify and analyze tiny DNA samples from a variety of sources, ranging from crime scenes to archaeological remains. For example, investigators have used PCR to analyze mitochondrial DNA obtained from the bones of Neandertals (see Chapter 22). Some of the Neandertal genome was sequenced in 2006, enabling researchers to compare it to equivalent parts of the human genome. A limitation of the PCR technique is that it is almost too sensitive. Even a tiny amount of contaminant DNA in a sample may become amplified if it includes a DNA sequence complementary to the primers, potentially leading to an erroneous conclusion. Researchers take appropriate precautions to avoid this and other technical pitfalls.

Review ■

■

■ ■

What are restriction enzymes? How are they used in recombinant DNA research? What are the relative merits of genomic DNA libraries, chromosome libraries, and cDNA libraries? How are DNA probes used? Why is the PCR technique valuable?

15.2 DNA ANALYSIS ■ ■ LEARNING OBJECTIVES 5 Distinguish among DNA, RNA, and protein blotting. 6 Describe the chain termination method of DNA sequencing.

DNA cloning in cells and the PCR technique have had major impacts on genetic analysis because they provide enough genetic material to address certain fundamental questions about specific genes and their RNA or protein products. For example, when— and how—is a gene expressed? Is a given gene identical from one individual to the next? Is it different from one species to the next? Where is the gene found within the genome? In this and the following section, we consider various techniques that help address these questions. We then examine some of the practical applications of DNA technology in the final section of the chapter.

Gel electrophoresis is used for separating macromolecules Mixtures of certain macromolecules—proteins, polypeptides, DNA fragments, or RNA—are separated by gel electrophoresis, a method that exploits the fact that these molecules carry charged groups that cause them to migrate in an electrical field. FIGURE 15-8 illustrates gel electrophoresis of DNA molecules. Nucleic acids migrate through the gel toward the positive pole of the electric field because they are negatively charged due to their phosphate groups. Because the gel slows down the large molecules more than the small molecules, the DNA is separated by size. Including DNA fragments of known size as standards ensures accurate measurements of the molecular weights of the unknown fragments. The

gel is stained after the DNA fragments are separated to show their number and location.

DNA, RNA, and protein blots detect specific fragments Geneticists can identify specific DNA fragments that hybridize with a complementary DNA probe. However, it is impossible to hybridize a probe to DNA fragments contained in a gel. For this reason the DNA is usually denatured and then transferred to a nitrocellulose or nylon membrane, which picks up the DNA like a blotter picks up ink. The resulting blot, which is essentially a replica of the gel, is incubated with a DNA probe, which hybridizes with any complementary DNA fragments. If the probe is radioactive, the blot is subjected to autoradiography. The resulting spots on the X-ray film correspond to the locations of the fragments in the gel that are complementary to the probe. Today, many probes are detected by chemical luminescence, which is analyzed by computer scanners, eliminating the need for autoradiography. This method of detecting DNA fragments—separating them by gel electrophoresis and then transferring them to a nitrocellulose or nylon membrane—is called a Southern blot, named for its inventor, Edward M. Southern of Edinburgh University in Scotland (FIG. 15-9). The procedure has widespread applications, such as being used to diagnose certain types of genetic disorders. For example, in some cases the DNA of a mutant allele is detected because it migrates differently in the gel than the DNA of its normal counterpart does. Similar blotting techniques are used to study RNA and proteins. When RNA molecules separated by electrophoresis are transferred to a membrane and detected using a nucleic acid probe, the result is called a Northern blot (as a pun on Southern’s name being associated with his technique). In the same spirit, the term Western blot is applied to a blot consisting of proteins or polypeptides previously separated by gel electrophoresis. (So far, no one has invented a type of blot that could be called an Eastern blot.) In the case of Western blotting, scientists recognize the polypeptides of interest by labeled antibody molecules that bind to them specifically. For example, Western blotting is used diagnostically to detect the presence of proteins specific to HIV-1, the AIDS virus.

Restriction fragment length polymorphisms are a measure of genetic relationships The variability of genes within a population can be studied in several ways. A traditional approach is based on the premise that no two individuals have the exact same number or position of restriction enzyme cutting sites in the genome: random DNA mutations and recombination may result in individuals with different lengths of fragments produced by a given restriction enzyme. Such restriction fragment length polymorphisms, commonly known as RFLPs, or “riflips,” can be used to determine how closely related different members of a population are. (The term polymorphism means “many forms” and refers to inherited differences found within a population.) A genetic polymorphism exists if individuals of two or more discrete types, or “morphs,” are found in a population.

RESEARCH METHOD Why Is It Used?

Gel electrophoresis separates molecules such as nucleic acids based on their size and charge as they migrate through an electrical field.

How Is It Done? 1

A researcher sets up an electrical field in a gel material, consisting of agarose or polyacrylamide, which is poured as a thin slab on a glass or Plexiglas holder. After the gel has solidified, the researcher loads samples containing a mixture of macromolecules (in this case, DNA fragments of different sizes) in wells formed at one end of the gel.

–

Standards of known sizes placed in well

2

When an electrical current is applied, the negatively charged DNA molecules move from the negative pole to the positive pole. The smallest DNA fragments travel the longest distance. The DNA bands are invisible at this point.

Anode –

Longer molecules

Direction of movement

Mixtures placed in well

Shorter molecules

Gel Buffer solution

+

+ Cathode

This gel contains separated DNA fragments. The gel is stained with ethidium bromide, a dye that binds to DNA and is fluorescent under UV light.

Eurelios/Photo Researchers, Inc.

3

FIGURE 15-8 Gel electrophoresis

Restriction enzymes are used to cut the DNA from two or more individuals, and the fragments are separated by gel electrophoresis, with the DNA from each individual in a separate lane; shorter fragments migrate farther than longer fragments. To visualize the fragments, researchers make a Southern blot of the DNA on the gel, which is then denatured and allowed to hybridize with a DNA probe. The resulting pattern of bands can then be compared.

RFLP analysis has been used when conducting paternity testing and when analyzing evidence found at crime scenes. RFLP analysis has also been used to help map the exact location of gene mutations, such as the mutation that causes cystic fibrosis. Although the RFLP technique has provided useful information in many areas of biology, it is rapidly being replaced by newer methods, such as automated DNA sequencing (discussed in the next section).

RESEARCH METHOD Why Is It Used?

The Southern blotting technique allows DNA separated by electrophoresis to be transferred to a filter in the same location as it was on the gel. It can then be hybridized with radioactive complementary probes.

Digest DNA with restriction enzymes.

1

How Is It Done?

DNA

2

Load DNA fragments on gel and separate by electrophoresis.

–

+

DNA fragments

Buffer solution 5

DNA fragments on filter are in same location as those on gel.

4

In blotting, buffer solution moves upward, transferring DNA fragments to a DNA-binding filter.

3

Agarose gel

Visualize DNA fragments using UV light.

Weight Longer DNA fragments

Absorbent paper Nitrocellulose filter Gel Wick Buffer

6

Place filter and radioactively labeled probe together in sealed bag so it can hybridize.

7

Shorter DNA fragments

Wash filter to remove excess probe and then expose filter to X-ray film; resulting autoradiograph shows hybridized DNA fragments.

Radioactive probe solution

FIGURE 15-9 The Southern blotting technique

Methods to rapidly sequence DNA have been developed One way to characterize DNA is to determine its sequence of nucleotides. Investigators use a cloned piece of DNA as a research tool for many different applications. For example, they may clone a gene to obtain the encoded protein for some industrial or medical process. Regardless of the particular application, researchers must know a great deal about the gene and how it functions. They usually start by determining the sequence of nucleotides.

In the 1990s, the advent of automated DNA-sequencing machines connected to powerful computers let scientists sequence huge amounts of DNA quickly and reliably. Automated DNA sequencing relies on the chain termination method that British biochemist Fred Sanger developed in 1974. In 1980, Sanger shared the Nobel Prize in Chemistry for this contribution. This was Sanger’s second Nobel Prize; his first was for his work on the structure of the protein insulin. Although DNA sequencing is now fully automated, many of the automated techniques are still based on Sanger’s chain termination method (FIG. 15-10).

RESEARCH METHOD The dideoxy chain termination method is a rapid, efficient way to sequence DNA.

How Is It Done?

Single-strand DNA fragment to be sequenced

5′ A

T

G

+ddATP 1

C

T

A

+ddCTP

T

G

C

T

+ddGTP

C

3′

C

+ddTTP

Four different reaction mixtures are used to sequence a DNA fragment; each contains a small amount of a single dideoxynucleotide, such as ddATP. Larger amounts of the four normal deoxynucleotides (dATP, dCTP, dGTP, and dTTP) plus DNA polymerase and radioactively labeled primers are also included.

A

Radioactive primer 5′ A T GC T A TGC TC C ddA C G A T A C G A G G 3′ ddA T A C G A G G

C

G

T

T A C G A T A C G A G G

3′ 5′

ddA C G A G G ddA G G

+ddATP

Direction of synthesis

5′

Reaction products from mixture containing dideoxyATP 2

3′

Larger fragments

Smaller fragments

A C G T

Courtesy of B. Slatko, New England Biolabs

Why Is It Used?

4 An exposed X-ray film of 3 The radioactive products of each reaction The random incorporation of dideoxyATP a DNA-sequencing gel. The (ddATP) into the growing chain generates mixture are separated by gel electrophoresis four lanes represent A, C, a series of smaller DNA fragments ending and located by exposing the gel to X-ray film. G, and T dideoxy reaction at all the possible positions where adenine The nucleotide sequence of the newly synthesized mixtures, respectively. is found in the newly synthesized fragments. DNA is read directly from the film (5′ → 3′ ). The sequence in the original template strand These correspond to positions where thymine is its complement (3′ → 5′ ). occurs in the original template strand.

FIGURE 15-10 The dideoxy chain termination method of DNA sequencing

The chain termination method of DNA sequencing is based on the fact that a replicating DNA strand that has incorporated a modified synthetic nucleotide, known as a dideoxynucleotide, cannot elongate beyond that point. Unlike a “normal” deoxynucleotide, which has a hydroxyl group on its 3¿ carbon, a dideoxynucleotide lacks a hydroxyl group on its 3¿ carbon. (Recall from Chapter 12 that a 3¿ hydroxyl group reacts each time a phosphodiester linkage is formed.) Thus, dideoxynucleotides terminate elongation during DNA replication. The researcher prepares four different reaction mixtures. Each contains multiple identical single-stranded copies of the DNA to be sequenced; DNA polymerase; appropriate radioactively la-

beled primers; and all four deoxynucleotides needed to synthesize DNA—dATP, dCTP, dGTP, and dTTP. Each mixture also includes a small amount of only one of the four dideoxynucleotides: ddATP, ddCTP, ddGTP, or ddTTP. The prefix “dd” refers to dideoxynucleotides, to distinguish them from deoxynucleotides, which are designated “d.” Here’s how the reaction proceeds in the mixture that includes ddATP. At each site where adenine is specified, occasionally a growing strand incorporates a ddATP, halting elongation so DNA fragments of varying lengths form in the reaction mixture. Each fragment that contains a ddATP marks a specific location where adenine would normally be found in the newly synthesized strand.

Similarly, in the reaction mixture that includes ddCTP, each fragment that contains ddCTP marks the position of a cytosine in the newly synthesized strand, and so on. The radioactive fragments from each reaction are denatured and then separated by gel electrophoresis, with each reaction mixture, corresponding to A, T, G, or C, occupying its own lane in the gel. The positions of the newly synthesized fragments in the gel can then be determined by autoradiography. Because the high resolution of the gel makes it possible to distinguish between fragments that differ in length by only a single nucleotide, the researcher can read off the sequence in the newly synthesized DNA one base at a time. The chain termination method is still used in many automated DNA-sequencing machines. However, the method of detection has changed in that the nucleotide sequence is no longer visualized by radioactivity. Instead, researchers label the primer or each of the four ddNTPs with a different-colored fluorescent dye. The computer uses a laser to read the fluorescence of the dye markers as the bases emerge from the end of a lane of electrophoresis medium (FIG. 15-11).

nary explosion of gene sequence data, largely due to automated sequencing methods. DNA sequence information is now kept in large computer databases, many of which are accessed through the Internet. Examples are databases maintained by the National Center for Biotechnology Information and by the Human Genome Organization (HUGO). Geneticists use these databases to compare newly discovered sequences with those already known to identify genes and to access many other kinds of information. By searching for DNA or amino acid sequences in a database, researchers can gain a great deal of insight into the function and structure of the gene product, the evolutionary relationships among genes, and the variability among gene sequences within a population. DNA sequencing of species has added much new information in support of evolution (see Chapter 18).

Review ■ ■

What is a Southern blot? What technique does automated DNA sequencing use? Outline the basic steps of this method.

Entire genomes have been sequenced using automated DNA sequencing

Sequencing machines today are very powerful and can decode about 1.5 million bases in a 24-hour period. These advances in sequencing technology have made it possible for researchers to study the nucleotide sequences of entire genomes in a wide variety of bacteria, archaea, and eukaryotic organisms. Much of this research received its initial impetus from the Human Genome Project, which began in 1990. The sequencing of the 3 billion base pairs of the human genome was essentially completed in 2001. The genomes of about 800 organisms have been sequenced now, and several hundred more are in various stages of planning or completion. We are in the middle of an extraordi-

15.3 GENOMICS ■ ■ LEARNING OBJECTIVES 7 8 9

Describe three important areas of research in genomics. Explain what a DNA microarray does and give an example of its research and medical potential. Define pharmacogenetics and proteomics.

© Mark Harmel/Alamy

Now that we have sequenced the genomes of humans and several hundred other species, what do we do with that information? Genomics is the emerging field of biology that studies the entire DNA sequence of an organism’s genome to identify all the genes, determine their RNA or protein products, and ascertain how the genes are regulated. As you will see, genomics has important practical applications in addition to answering scientific questions. The science of genomics has several research areas, including structural genomics, functional genomics, comparative genomics, and metagenomics. Structural genomics is concerned with mapping and sequencing genomes. Functional genomics deals with the functions of genes and nongene sequences in genomes. Comparative genomics involves comparing the genomes of different species to determine what changes in DNA have occurred during the course of evolution. This knowledge furthers our understanding of evolutionary relationships because genes with little or no variation among species are functionally important in the species in which they are found. FIGURE 15-11 Animated A small segment of the printout of automated Metagenomics uses the techniques of genomics to DNA-sequencing results analyze communities of microorganisms instead The computer produces the series of peaks shown as it reads the fluorescing bands on the of individual organisms. Metagenomics enables reelectrophoresis gel. The nucleotide base adenine is shown in green, guanine in black, cytosine searchers to identify and study all the genes in a comin blue, and thymine in red. The DNA sequence assigned by the computer appears at the top of the printout. munity at the same time.

Identifying protein-coding genes is useful for research and for medical applications The majority of the human genome contains DNA sequences that do not code for proteins. Although the entire genome is of interest to biologists, the protein-coding regions are of particular interest to medical researchers. Scientists have compiled tens of thousands of short (25 to 30 nucleotides) cDNA sequences, known as expressed sequence tags (ESTs), that can help identify proteincoding genes. A large data set containing thousands of human ESTs is available to scientists via the Internet. Using gene-hunting software, biologists working on a particular DNA sequence can access the data set and match their DNA sequence to a known EST sequence. Many ESTs in the data set have been identified by comparing them to known DNA sequences from other species. For example, the human telomerase gene TRT was identified by matching its human EST sequence to that of a known yeast telomerase sequence. (Recall from Chapter 12 that telomerase adds repetitive nucleotide sequences to the ends, or telomeres, of eukaryotic chromosomes.)

One way to study gene function is to silence genes one at a time RNA interference (RNAi), first discovered in the mid-1990s, can be used to quickly determine the function of a specific gene. You may recall from Chapter 13 that RNAi is caused by small RNA molecules that interfere with the expression of genes or, more precisely, their mature mRNA transcripts. RNA interference involves small interfering RNAs, microRNAs, and a few other kinds of short RNA molecules. Biologists hypothesize that RNAi, which occurs in many eukaryotic organisms, originally evolved in nature because it protected cells against viruses that have double-stranded RNA molecules. After a protein-coding gene is identified, perhaps by using ESTs, the function of that gene can be studied by using RNA interference to inactivate the gene. To do this, biologists synthesize a short stretch of RNA that is complementary to part of the DNA sequence of the gene being examined. The RNA is put into cells, and after the gene is silenced, biologists observe any changes in the phenotype to help determine the function of the missing protein. Gene targeting also reveals gene function Another research tool that reveals gene function is gene targeting, a procedure in which the researcher chooses and “knocks out,” or inactivates, a single gene in an organism. Knockout mice have been particularly useful in studying the roles of orthologous genes with unknown functions in mammals, including humans. Orthologous genes are homologous genes found in different species because they were inherited from a common ancestor. The roles of the inactivated gene are determined by observing the phenotype of the mice bearing the knockout gene. If the gene codes for a protein, for example, studying individuals who lack that protein helps the researcher identify the function of the protein. Because 98% to 99% of the loci of mice are orthologous to human loci, information about knockout genes in mice also provides details about human genes. Unlike RNA interference, gene targeting is a lengthy procedure. It takes about a year for scientists to develop a new strain of knockout

mice through gene targeting. A nonfunctional, or knockout, gene is cloned and introduced into mouse embryonic stem cells (ES cells) (also see Chapters 1 and 17). Once inside the ES cells, genes are relatively easy to handle because, like cancer cells, ES cells grow in culture indefinitely. Most important, if placed in a mouse embryo, ES cells divide and produce all the cell types normally found in the mouse. In a tiny fraction of these ES cells, the introduced knockout gene becomes physically associated with the corresponding gene in a chromosome. If this occurs, the gene on the chromosome and the knockout gene tend to exchange DNA segments in a process called homologous recombination. In this way, the knockout allele replaces the normal allele in the mouse chromosome. Researchers inject into early mouse embryos ES cells they think are likely to carry a knockout gene and let the mice develop to maturity. The researchers generally study animals that carry the knockout gene in every cell. However, because many genes are essential to life, researchers have modified the knockout technique to develop strains in which a specific gene is selectively inactivated in only one cell type. Research laboratories around the world have developed thousands of different strains of knockout mice, each displaying its own characteristic phenotype, and the number continues to grow. Gene targeting in mice is providing answers to basic biological questions relating to the development of embryos, the development of the nervous system, and the normal functioning of the immune system. This technique has great potential for revealing more about various human diseases, especially because many diseases have a genetic component. Geneticists are using gene targeting to study cancer, heart disease, sickle cell anemia, respiratory diseases such as cystic fibrosis, and other disorders. Mutagenesis screening reveals the genes involved in a particular phenotype Many large-scale mutagenesis screening projects in mice and other organisms are currently underway. In mutagenesis screening, researchers treat male mice with chemical mutagens that cause mutations in DNA. They then breed the males with normal females and screen their offspring for unusual phenotypes. Unlike gene targeting, mutagenesis does not necessarily disable a gene completely; instead, it usually causes small, random mutations that change the properties of the proteins that the DNA encodes. The scale of mutagenesis screening is enormous; one screening project in Germany has screened some 28,000 different mouse mutants for changes in phenotypes.

DNA microarrays are a powerful tool for studying gene expression DNA microarrays provide a way to study patterns of gene expression in a variety of organisms. Each spot in a DNA microarray consists of a single-stranded probe placed on a glass slide or chip (FIG. 15-12). First, a mechanical robot prepares the microarray. It spots each location on the grid with thousands to millions of copies of a specific complementary DNA (cDNA) strand. Each spot, known as a microdot, contains many copies of a specific singlestranded cDNA molecule. These were synthesized to make a complementary copy of a specific mRNA, which was then amplified by PCR.

RESEARCH METHOD Why Is It Used?

DNA microarray analysis indicates gene expression levels of hundreds or even thousands of specific genes in different cell types. This technique helps scientists understand which genes are active (or inactive) in untreated (control) cells as well as in cells treated with a specific drug.

How Is It Done? 1

Prepare microarray. Each microdot contains multiple copies of a specific single-stranded probe (such as cDNA).

Treated cell

Untreated (control) cell 2

Mature mRNA

Prepare cDNA from two cell populations (treated and control).

Mature mRNA

Reverse transcriptase

cDNA copy of mRNA 3

cDNA

cDNA copy of mRNA

Tag each cDNA with different fluorescent dye.

mRNA (discard)

cDNA

mRNA (discard)

4

5

Laser 1 Laser 2

Reverse transcriptase

Hybridize two cDNA populations to array.

Scan array to identify fluorescence where hybridization has occurred.

Gene in treated cell that increased activity, compared to control

Emissions 6

FIGURE 15-12 DNA microarray analysis

Computer analysis produces color-coded readout.

Gene in treated cell that decreased activity, compared to control Gene that was active in both treated and untreated cells Gene that was inactive in both treated and untreated cells

Researchers then isolate mature mRNA molecules from two cell populations—for example, liver cells treated with a newly developed drug and control liver cells not treated with the drug. Liver cells are often tested because the liver produces many enzymes that metabolize foreign molecules, including drug molecules. Drugs the liver cannot metabolize, or metabolizes weakly, may be too toxic to be used. Researchers use the isolated mRNA molecules to prepare cDNA molecules for each cell population, which are then tagged with different-colored fluorescent dyes. For example, the treated cells’ cDNA molecules might be labeled with red dye and the untreated cells’ cDNA molecules with green dye. Then researchers add the two cDNA populations to the array, and some of the cDNA subsequently hybridizes (forms base pairs with) the cDNA on the array. After washing it to remove any cDNA that has not hybridized to the array, investigators scan the array with lasers to identify red and green fluorescence where hybridization has occurred. Computer analysis of the ratio of red to green fluorescence at each spot in the array produces a color-coded readout that researchers can further analyze. For example, a medical researcher might compare the overall pattern of gene activity in cells treated with an experimental drug with that of cells exposed to known drugs and toxins. If the pattern of gene activity for the treated cells matches that of cells treated with a toxin that damages the liver, the drug will probably not go into clinical trials. DNA microarrays enable researchers to compare the activities of thousands of genes in cell types from various tissue samples and in normal and diseased cells. Because cancer and other diseases exhibit altered patterns of gene expression, DNA microarrays have the potential to identify disease-causing genes or the proteins they code for, which can then be targeted by therapeutic drugs. Here’s an example of the application. Diffuse large B-cell lymphoma is a cancer of white blood cells that has different subtypes. In 2002, researchers used microarrays to identify 17 genes that are active in various combinations for each subtype. Using DNA microarrays, researchers determined which combination of those 17 genes is active in each subtype of this form of cancer. Armed with the knowledge of which cancer subtype a patient has, physicians can choose the treatment that will probably be most effective for that patient. Examining patterns of gene activity with DNA microarrays also helps medical scientists identify which patients will probably remain free of cancer after treatment and which will probably relapse.

The Human Genome Project stimulated studies of genome sequencing for other species To aid in analyzing the human genome, investigators carried out comparative genomics by sequencing and mapping studies simultaneously on the mouse and rat genomes. Mice and rats were obvious choices for DNA sequencing because they have been studied for almost a century and much is known about their biology. Mice and rats are sufficiently different from humans that any conserved (that is, identical) sequences found in both rodents and humans are probably functionally important. In addition, mice are similar enough to humans that both share many physiological traits, including some of the same diseases. To demonstrate the shared similarity between the mouse and human genomes, a consortium of scientists published a study in 2002 that

compared the mouse chromosome 16 with the human genome. Only 14 of the 731 known genes on the mouse chromosome had no human counterpart; the remaining 717 genes appeared in one form or another in the human genome. Comparison of the DNA sequences and chromosome organization of related genes from different species is a powerful tool for identifying the elements essential for their functions. If a human gene has an unknown function, researchers can often deduce clues about its role by studying the equivalent gene in another species, such as a mouse or rat. Investigators have also sequenced the genomes of other model vertebrate organisms, such as the zebrafish and pufferfish. Biologists have used the zebrafish for many years as a model of vertebrate development because its embryonic stages are transparent and therefore convenient to examine. The pufferfish has the smallest known genome of all vertebrates. By comparing the human genome to that of the pufferfish, researchers have identified shared sequences that have not changed in several hundred million years. Because these sequences have resisted the selective forces of evolution, they may be genes or regulatory elements that direct vertebrate development and are essential in all vertebrates. Genome analysis of commercially important organisms, such as salmon, chickens, pigs, and rice, is also a priority. For example, scientists have published the rice genome, the genome of the parasite that causes malaria, and the genome of the parasite that causes African sleeping sickness. One of the most significant genome-sequencing efforts from an evolutionary perspective focuses on nonhuman primates, such as the chimpanzee (whose sequence data were published in 2005). Comparing the DNA sequences of humans and our closest living relatives is helping biologists understand the genetic changes that occurred during human evolution, including which genes govern mental and linguistic capabilities.

Several scientific fields—bioinformatics, pharmacogenetics, and proteomics— have emerged We have already mentioned that the amount of genomic data now available requires that it be evaluated using powerful computers. The discipline known as bioinformatics, or biological computing, includes the storage, retrieval, and comparison of nucleotide or amino acid sequences within a given species and among different species. Bioinformatics uses powerful computers and sophisticated software to store, manage, and analyze large amounts of data generated by sequencing and other technologies. For example, as researchers determine new DNA sequences, automated computer programs scan the sequences for patterns typically found in genes. Through comparison of the databases of DNA sequences from different organisms, bioinformatics has already led to insights about gene identification, gene function, and evolutionary relationships. The new science of gene-based medicine known as pharmacogenetics studies how genetic variation among patients affects the action of drugs in individuals. The goal of pharmacogenetics is to customize drugs to match an individual’s genetic makeup. Currently, a physician does not know in advance whether a particular medication will benefit a patient or cause severe side effects. An individual’s genes, especially those that code for drug-metabolizing

enzymes, largely determine that person’s response to a specific drug. Pharmacogenetics takes into account the subtle genetic differences among individuals. In as few as 5 to 10 years, patients may take routine genetic screening tests before a physician prescribes a drug. Pharmacogenetics, like other fields in human genetics, presents difficult ethical questions. The genetic testing that will become an everyday part of pharmacogenetics raises issues of privacy, genetic bias, and potential discrimination. We consider these issues in Chapter 16. Individualized genetic testing may cause people to worry unnecessarily about a genetic disease for which they may never develop symptoms. Most common diseases result from a complex interplay between multiple genes and nongenetic, or environmental, factors. (Recall from Chapter 11 that the environment is an important factor influencing gene expression.) In humans, healthy environmental factors include proper diet, adequate exercise, and not smoking. The study of all the proteins expressed by a cell at a given time is known as proteomics. For example, scientists have identified all the proteins made by a given kind of cell, but the process is much more complicated than sequencing the human genome. Some genes, for example, encode several different proteins. (See Figure 14-14 for an example of alternative splicing.) Also, every somatic cell in the human body has essentially the same genome, but cells in different tissues vary greatly in the kinds of proteins they produce (recall the earlier discussion of DNA microarrays). Protein expression patterns vary not only in different tissues but also at different stages in the development of a single cell. Scientists want to understand the role of each protein in a cell, the way the various proteins interact, the 3-D structure for each protein, how proteins are spatially arranged within the cell’s cytosol and organelles, and protein profiles in diseased cells. While advancing biological knowledge, these goals also promise advances in medicine. If they know the shape of a protein associated with a type of cancer or other disease, scientists may be able to develop drugs that bind to active sites on that protein, turning its activity off. Today, the pharmaceutical industry has drugs that target about 500 proteins in the cell. However, researchers estimate proteomics may yield 10,000 to 20,000 additional protein targets.

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How would you distinguish among structural, functional, and comparative genomics? What is a DNA microarray? What is pharmacogenetics? proteomics?

15.4 APPLICATIONS OF DNA TECHNOLOGIES ■ ■ LEARNING OBJECTIVE 10 Describe at least one important application of recombinant DNA technology in each of the following fields: medicine, DNA fingerprinting, and transgenic organisms.

Recombinant DNA technology has provided both a new and unique set of tools for examining fundamental questions about cells and new approaches to applied problems in many other fields.

In some areas, the production of genetically modified proteins and organisms has begun to have considerable impact on our lives. The most striking of these has been in medicine.

DNA technology has revolutionized medicine In increasing numbers of cases, doctors perform genetic tests to determine whether an individual has a particular genetic mutation associated with such disorders as Huntington’s disease, hemophilia, cystic fibrosis, Tay-Sachs disease, breast cancer, and sickle cell anemia. Gene therapy, the use of specific DNA to treat a genetic disease by correcting the genetic problem, is another application of DNA technology that is currently in its infancy. Because genetic testing and gene therapy focus almost exclusively on humans, these applications of DNA technology are discussed in Chapter 16. We focus our discussion here on the use of DNA technology to produce medical products. Human insulin produced by E. coli became one of the first genetically modified proteins approved for use by humans. Before the use of recombinant DNA techniques to generate genetically altered bacteria capable of producing the human hormone, insulin was derived exclusively from other animals. Many diabetic patients become allergic to the insulin from animal sources because its amino acid sequence differs slightly from that of human insulin. The ability to produce the human hormone by recombinant DNA methods has resulted in significant medical benefits to insulindependent diabetics. Genetically modified human growth hormone (GH) is available to children who need it to overcome growth deficiencies, specifically pituitary dwarfism (see Chapter 49). In the past, human GH was obtainable only from cadavers. Only small amounts were available, and evidence suggested that some of the preparations from human cadavers were contaminated with infectious agents similar to those that cause bovine spongiform encephalopathy, or mad cow disease (discussed in Chapter 24). The list of products produced by molecular modification is continually growing. For example, tissue plasminogen activator (TPA), a protein that prevents or dissolves blood clots, is used to treat cardiovascular disease. If administered shortly after a heart attack, TPA reduces the risk of a subsequent attack. Tissue growth factor-beta (TGF-b) promotes the growth of blood vessels and skin, and is used in wound and burn healing. Researchers also use TGF-b in tissue engineering, a developing technology to meet the pressing need for human tissues and, eventually, organs for transplantation by growing them from cell cultures. The U.S. Food and Drug Administration (FDA) has approved tissue-engineered skin grafts for individuals who have been badly burned, and tissue-engineered cartilage is used for joint repairs. Researchers are investigating the development and use of cardiac cells from ES cells; these cells could be used to repair the heart following heart attacks. Work is also being done on building organs such as bladders and hearts. Hemophilia A is treated with human clotting factor VIII. Before the development of recombinant DNA techniques, factor VIII was available only from human or animal-derived blood, which posed a risk of transmitting infectious agents, such as HIV. Dornase Alpha (DNase) improves respiratory function (it helps to liquefy and thin sticky mucus) in people with cystic fibrosis.

DNA fingerprinting has numerous applications The analysis of DNA fragments extracted from an individual, which is unique to that individual, is known as DNA fingerprinting. DNA fingerprinting has many applications in humans and other organisms, as in the following examples: 1. Analyzing evidence found at crime scenes (forensic analysis) (FIG. 15-13) 2. Identifying mass disaster victims 3. Proving parentage in dogs for pedigree registration purposes 4. Identifying human cancer cell lines 5. Studying endangered species in conservation biology 6. Tracking tainted foods 7. Studying the genetic ancestry of human populations 8. Clarifying disputed parentage 9. Exonerating prisoners wrongfully convicted of a crime Historically, DNA fingerprinting relied on RFLPs (discussed earlier). Today, many techniques for DNA fingerprinting rely on PCR amplification, restriction enzyme digestion, and Southern blot hybridization to detect molecular markers. The most useful molecular markers are highly polymorphic within the human population (recall our earlier discussion of polymorphism). Short tandem repeats (STRs) are molecular markers that are short sequences of repetitive DNA—up to 200 nucleotide bases with a simple repeat pattern such as GTGTGTGTGT or CAGCAGCAGCAG. Because they vary in length from one individual to another, STRs are highly polymorphic. This characteristic makes them useful with a high degree of certainty in identifying individuals. If enough markers are compared, the odds that two people taken at random from the general population would have identical DNA profiles may be as low as one in several billion. The FBI uses a set of STRs from 13 different markers to establish a unique DNA profile for an individual. Such a profile distinguishes that person from every other individual in the United States, except an identical twin. Recall that DNA is also found in the cell’s mitochondria. Whereas nuclear DNA occurs in two copies per cell—one on each of the homologous chromosomes—mitochondrial DNA has as

Cellmark Diagnostics, Abingdon, U.K.

Recombinant DNA technology is increasingly used to produce vaccines that provide safe and effective immunity against infectious diseases in humans and animals. One way to develop a recombinant vaccine is to clone a gene for a surface protein produced by the disease-causing agent, or pathogen; the researcher then introduces the gene into a nondisease-causing vector. The vaccine, when delivered into the human or animal host, stimulates an immune response to the surface-exposed protein. As a result, if the pathogen carrying that specific surface protein is encountered, the immune system targets it for destruction. Human examples of antiviral recombinant vaccines include vaccines for influenza A, hepatitis B, and polio. Recombinant vaccines are also being developed against certain bacterial diseases and human cancers. For example, in 2006 the Food and Drug Administration approved a recombinant vaccine against several types of human papillomavirus (HPV). The vaccine, which is administered to females ages 9 to 26, protects against the 70% of cervical cancers caused by HPV.

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FIGURE 15-13 Animated DNA fingerprinting These DNA fingerprints show DNA from a crime scene (in middle), along with the DNA profiles of seven suspects. Can you pick the suspect whose DNA profile matches blood from the crime scene?

many as 100,000 copies per cell. Therefore, mitochondrial DNA is the molecule of choice for DNA fingerprinting in which biological samples have been damaged—identifying exhumed human remains, for example. DNA fingerprinting has revolutionized law enforcement. In 1990, the FBI established the Combined DNA Index System (CODIS), consisting of DNA databases from all 50 states. A DNA profile of an unknown suspect can be compared to millions of DNA profiles of convicted offenders in the database, often resulting in identification of the suspect. The DNA from the unknown suspect may come from blood, semen, bones, teeth, hair, saliva, urine, or feces left at the crime scene. Tiny amounts of human DNA have even been extracted from cigarette butts, licked envelopes or postage stamps, dandruff, fingerprints, razor blades, chewing gum, wristwatches, earwax, debris from under fingernails, and toothbrushes. If applied properly, DNA fingerprinting has the power to identify the guilty and exonerate the innocent. Hundreds of convicted individuals have won new trials and have been subsequently released from incarceration based on correlating DNA profiles with physical evidence from the crime scene. One limitation arises from the fact that the DNA samples are usually small and may have been degraded. Obviously, great care must be taken to prevent contamination of the samples. This is especially crucial if the PCR technique is used to amplify DNA.

Transgenic organisms have foreign DNA incorporated into their cells Plants and animals in which foreign genes have been incorporated are called transgenic organisms. Researchers use varied approaches to insert foreign genes into plant or animal cells. They often use viruses as vectors, although other methods, such as the direct injection of DNA into cells, have also been applied.

FIGURE 15-14 A transgenic mouse The mouse on the right is normal, whereas the mouse on the left is a transgenic animal that expresses rat growth hormone in large amounts.

whose synthesis is stimulated by the presence of toxic amounts of heavy metals such as zinc. The researchers used metallothionein regulatory sequences as a switch to turn the production of rat GH on and off at will. After injecting the modified gene into mouse embryo cells, they implanted the embryos into the uterus of a mouse and let them develop. Because of the difficulty in manipulating the embryos without damaging them, the gene transplant succeeded in only a small percentage of the animals. When exposed to small amounts of zinc, these transgenic mice produced large amounts of growth hormone because the liver is a much larger organ than the pituitary gland. The mice grew rapidly, and one mouse, which developed from an embryo that had received two copies of the GH gene, grew to more than double the normal size (FIG. 15-14). As might be expected, such mice often transmit their increased growth capability to their offspring. Transgenic animals can produce genetically modified proteins Certain transgenic animal strains produce milk containing foreign proteins of therapeutic or commercial importance. For example, researchers have introduced into sheep the gene for the human protein tissue plasminogen activator (TPA). Producing transgenic livestock (such as pigs, sheep, cows, and goats) that secrete foreign proteins in their milk is known informally as “pharming,” a combination of “pharmaceuticals” and “farming” (FIG. 15-15). In pharming, recombinant genes are fused to the regulatory sequences of the milk protein genes, and such genes are therefore activated only in mammary tissues involved in milk production. The advantages of obtaining a protein drug from milk are that

Pentti Vänskä, Helsingin Sanomat International Edition

R. L. Brinster, University of Pennsylvania Medical School

Transgenic animals are valuable in research Transgenic animals are usually produced by injecting the DNA of a particular gene into the nucleus of a fertilized egg cell or of ES cells. The researcher then implants the eggs into the uterus of a female and lets them develop. Alternatively, the researcher injects genetically modified ES cells into isolated blastocysts, an early stage in embryonic development, and then implants them into a foster mother. Injecting DNA into cells is not the only way to produce transgenic animals. Researchers may use viruses as recombinant DNA vectors. RNA viruses called retroviruses make DNA copies of themselves by reverse transcription. Sometimes the DNA copies become integrated into the host chromosomes, where they are replicated along with host DNA. Transgenic animals provide valuable applications over a wide range of research areas, such as regulation of gene expression, immune system function, genetic diseases, viral diseases, and genes involved in the development of cancer. The laboratory mouse is a particularly important model organism for these studies. In a classic study of the control of gene expression, University of Pennsylvania geneticist Ralph L. Brinster in 1983 produced transgenic mice carrying a gene for rat growth hormone (see Fig. 17-18). Brinster and his coworkers wanted to understand the controls that let certain genes be expressed in some tissues and not in others. The pituitary gland of a mouse normally produces small amounts of GH, and these researchers reasoned that other tissues might also be capable of producing the hormone. First they isolated the GH gene from a library of genomic rat DNA. They combined it with the promoter region of a mouse gene that normally produces metallothionein, a protein that is active in the liver and

FIGURE 15-15 Transgenic “pharm” cows These cows contain a human gene that codes for lactoferrin, a protein found in human mothers’ milk and in secretions such as tears, saliva, bile, and pancreatic fluids. Lactoferrin is one of the immune system’s lines of defense against disease-causing organisms. The cows secrete human lactoferrin in their milk.

Transgenic plants are increasingly important in agriculture People have selectively bred plants for thousands of years. The success of such efforts depends on desirable traits in the variety of plant selected or in closely related wild or domesticated plants whose traits are transferred by crossbreeding. Local varieties or closely related species of cultivated plants often have traits, such as disease resistance, that agriculturalists could advantageously introduce into varieties more useful to modern human needs. If genes are introduced into plants from strains or species with which they do not ordinarily interbreed, the possibilities for improvement increase greatly. The most widely used vector system for introducing recombinant genes into many types of plant cells is the crown gall bacterium, Agrobacterium tumefaciens. This bacterium normally produces plant tumors by introducing a plasmid, called the Ti plasmid, into the cells of its host; Ti stands for “tumor-inducing.” The Ti plasmid induces abnormal growth by forcing the plant cells to produce elevated levels of a plant growth hormone called cytokinin (discussed in Chapter 38). Geneticists “disarm” the Ti plasmid so that it does not induce tumor formation and then use it as a vector to insert genes into plant cells. The cells into which the altered plasmid is introduced are essentially normal except for the inserted genes. Genes placed in the plant genome in this fashion may be transmitted sexually, via seeds, to the next generation, but they can also be propagated asexually, for example, by taking cuttings. Unfortunately, not all plants take up DNA readily, particularly the cereal grains that are a major food source for humans. One useful approach has been the development of a genetic “shotgun.” Researchers coat microscopic gold or tungsten fragments with DNA and then shoot them into plant cells, penetrating the cell walls. Some of the cells retain the DNA, which transforms them. Those cells can then be cultured and used to regenerate an entire plant (see Fig. 17-2). Geneticists have successfully used such an approach to transfer a gene for resistance to a bacterial disease into cultivated rice from one of its wild relatives. An additional complication of plant genetic modification is that about 120 plant genes lie in the DNA of the chloroplasts; the other 3000 or so genes that plants require to function are in the nucleus. Chloroplasts are essential in photosynthesis, which is the basis for plant productivity. Because great agricultural potential hinges on improving the productivity of photosynthesis, developing methods for changing the part of the plant’s DNA within the chloroplast is desirable. Dozens of labs are currently studying methods of chloroplast engineering, although progress has been slow. In 2001, Australian plant physiologists reported that they had altered rubisco, the key carbon-fixing enzyme of photosynthesis (see Chapter 9), by changing one of the genes in the chloroplast genome. The United States is the world’s top producer of transgenic crops, also known as genetically modified (GM) crops. In 2007,

U.S. farmers planted 57.7 hectares (143 million acres) of GM crops. Globally, 51% of the soybean crop, 31% of corn, 13% of cotton, and 5% of canola are GM crops. Selected Applications of Transgenic Plants Agricultural geneticists are developing GM plants that are resistant to insect pests, viral and fungal diseases, heat, cold, herbicides, salty or acidic soil, and drought. For example, genetically modified papaya trees, which are resistant to the ringspot virus, have been grown in Hawaii for more than 10 years. A variety of corn that has been genetically modified to be resistant to drought is currently being tested in field trials (FIG. 15-16).

Courtesy of the Monsanto Corporation

potentially it can be produced in large quantities and that it can be harvested simply by milking the animal. The protein is then purified from the milk. The introduction of the gene does not harm the animals, and because the offspring of the transgenic animal usually produce the recombinant protein, transgenic strains are established.

(a) Note the poor yield in genetically unmodified corn plants used as a control.

(b) Genetically modified corn plants withstood drought better than the unmodified corn.

FIGURE 15-16 Animated Field test of transgenic corn with resistance to drought

Corn has also been genetically modified to contain the Bt gene, a bacterial gene that codes for a protein with insecticidal properties; Bt stands for the bacterium’s scientific name, Bacillus thuringiensis. Bt corn, introduced in the United States in 1996, does not need periodic sprays of chemical insecticides to control the European corn borer, the most damaging insect pest of corn in the United States and Canada. DNA technology also has the potential to develop crops that are more nutritious. For example, in the 1990s geneticists modified rice to produce high quantities of b-carotene, which the human body uses to make vitamin A. In developing countries, vitamin A deficiency is a leading cause of irreversible blindness in children. Vitamin A deficiency also makes children more susceptible to measles and other infectious diseases. Because rice is the staple diet in many countries with vitamin A deficiency, the widespread use of GM rice with b-carotene could help prevent blindness in many of the world’s children. However, GM rice is not yet approved for human consumption. Like some transgenic animals, certain transgenic plants can potentially be “pharmed” to produce large quantities of medically important proteins. Examples include antibodies against the herpes virus and a patient-specific vaccine for lymphoma, a type of cancer. Some people are concerned about the health effects of consuming foods derived from GM crops and think that such foods should be restricted. For example, critics say that some consumers may develop food allergies. Scientists also recognize this concern and routinely screen new GM crops for allergenicity. There is also an ongoing controversy as to whether GM foods should be labeled. The FDA and most scientists think such labeling would be counterproductive because it would increase public anxiety over a technology that is as safe as traditional breeding methods. In 1996, the U.S. Court of Appeals upheld the FDA position that labeling should not be required.

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In what way has the production of human insulin by recombinant DNA methods had significant medical advantages for diabetics? What are short tandem repeats (STRs), and why are they so useful in DNA fingerprinting? Why do gene targeting and mutagenesis screening in mice have potential benefits for humans?

15.5 DNA TECHNOLOGY HAS RAISED SAFETY CONCERNS ■ ■ LEARNING OBJECTIVE 11 Describe at least two safety issues associated with recombinant DNA technology and explain how these issues are being addressed.

When recombinant DNA technology was introduced in the early 1970s, many scientists considered the potential misuses at least as significant as the possible benefits. The possibility that an organism with undesirable environmental effects might be accidentally produced was of great concern. Scientists feared that new strains

of bacteria or other organisms, with which the world has no previous experience, might be difficult to control. The scientists who developed the recombinant DNA methods insisted on stringent guidelines for making the new technology safe. Experiments in thousands of university and industry laboratories over more than 35 years have shown that recombinant DNA manipulations can be carried out safely. Geneticists have produced strains of E. coli that cannot survive outside of the laboratory. Researchers carry out experiments that might present unusual risks in facilities designed to hold pathogenic organisms; this precaution ensures that researchers can work with them safely. So far, no evidence suggests that researchers have accidentally cloned hazardous genes or have released dangerous organisms into the environment. However, malicious intentional manipulation of dangerous genes certainly remains a concern. As the safety of the experiments has been established, scientists have relaxed many of the restrictive guidelines for using recombinant DNA. Stringent restrictions still exist, however, in certain areas of recombinant DNA research where there are known dangers or where questions about possible effects on the environment are still unanswered. These restrictions are most evident in research that proposes to introduce transgenic organisms into the wild, such as agricultural strains of plants whose seeds or pollen might spread in an uncontrolled manner. A great deal of research now focuses on determining the effects of introducing transgenic organisms into a natural environment. Carefully conducted tests have shown that transgenic organisms are not dangerous to the environment simply because they are transgenic. It is, however, important to assess the risks of each new recombinant organism. Scientists determine whether the organism has characteristics that might cause it to be environmentally hazardous under certain conditions. For example, if geneticists have modified a transgenic crop plant to resist an herbicide, could that gene be transferred, via pollen or some other route, to that plant’s weedy relatives, generating herbicide-resistant “superweeds”? In 2003, ecologists at the University of Tennessee, Knoxville, announced the crossing of transgenic oilseed rape plants that contained the Bt gene with their wild relative, which is a weed. They crossed the resulting hybrids with the wild relative again and then tested its ability to compete with other weeds in a field of wheat. The transgenic weed was a poor competitor and had less effect on wheat production than its wild relatives in a control field. These results, although encouraging, must be interpreted with care. Scientists must evaluate each transgenic crop plant individually to see if there is gene flow to wild relatives, and if so, what effects might result. Other concerns relate to plants modified to produce pesticides, such as the Bt toxin. The future of the Bt toxin in transgenic crops is not secure because low levels of the insecticide could potentially provide ideal conditions for selection for resistant individuals in the insect population. It appears certain insects may evolve genetic resistance to the Bt toxin in transgenic plants in the same way they evolve genetic resistance to chemical insecticides. Another concern is that nonpest species could be harmed. For example, people paid a great deal of attention to the finding that monarch butterfly larvae raised in the laboratory are harmed if they are fed pollen from Bt corn plants. However, more recent

studies suggest that monarch larvae living in a natural environment do not consume enough pollen to cause damage. Also, evidence indicates that the harm to nonpest species from spraying pesticides is far greater than any harm from transgenic crops. Environmental concerns about transgenic animals also exist. Several countries are in the process of developing fast-growing transgenic fish, usually by inserting a gene that codes for a growth hormone. Transgenic Atlantic salmon, for example, grow up to six times as fast as nontransgenic salmon grown for human consumption. The transgenic fish do not grow larger than other fish, just faster. The benefits of such genetically enhanced fish include reduced pressure on wild fisheries and less pollution from fish farms. However, if the transgenic fish escaped from the fish farm, what effect would they have on wild relatives? To address this concern, the researchers are developing only nonreproducing female transgenic salmon.

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To summarize, DNA technology in agriculture offers many potential benefits, including higher yields by providing disease resistance, more nutritious foods, and the reduced use of chemical pesticides. However, like other kinds of technology, molecular modification poses some risks, such as the risk that genetically modified plants and animals could pass their foreign genes to wild relatives, causing unknown environmental problems. The science of risk assessment, which uses statistical methods to quantify risks so they can be compared and contrasted, will help society decide whether to ignore, reduce, or eliminate specific risks of molecularly modified organisms.

Review What are some of the environmental concerns regarding transgenic organisms? What kinds of information does society need to determine if these concerns are valid?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

15.1 (page 324) 1 Explain how a typical restriction enzyme cuts DNA molecules and give examples of the ways in which these enzymes are used in recombinant DNA technology. ■ Recombinant DNA technology isolates and amplifies specific sequences of DNA by incorporating them into vector DNA molecules. Researchers then clone—propagate and amplify—the resulting recombinant DNA in organisms such as E. coli. ■ Researchers use restriction enzymes to cut DNA into specific fragments. Each type of restriction enzyme recognizes and cuts DNA at a highly specific base sequence. Many restriction enzymes cleave DNA sequences to produce complementary, single-stranded sticky ends. Learn more about restriction enzymes by clicking on the figure in CengageNOW. ■

Genes present in genomic DNA and chromosome libraries from eukaryotes contain introns, regions that do not code for protein. Those genes can be amplified in bacteria, but the protein is not properly expressed. Because introns have been removed from mature mRNA molecules, eukaryotic genes in cDNA libraries can be expressed in bacteria to produce functional protein products. 3 Explain how researchers use a DNA probe. ■ Researchers use a radioactive DNA sequence as a DNA probe to screen thousands of recombinant DNA molecules in bacterial cells to find the colony that contains the DNA of interest. ■

Watch the process for using a DNA probe to find bacterial cells with a specific recombinant DNA molecule by clicking on the figure in CengageNOW.

4 Describe how the polymerase chain reaction amplifies DNA in vitro.

Geneticists often construct recombinant DNA molecules by allowing the ends of a DNA fragment and a vector, both cut with the same restriction enzyme, to associate by complementary base pairing. Then DNA ligase covalently links the DNA strands to form a stable recombinant molecule. Learn more about the formation of recombinant DNA by clicking on the figure in CengageNOW.

2 Distinguish among a genomic DNA library, a chromosome library, and a complementary DNA (cDNA) library; explain why one would clone the same eukaryotic gene from both a genomic DNA library and a cDNA library. ■ A genomic DNA library contains thousands of DNA fragments that represent the total DNA of an organism. A chromosome library contains all the DNA fragments from a specific chromosome. Each DNA fragment of a genomic DNA or chromosome library is stored in a specific bacterial strain. ■ A cDNA library is produced using reverse transcriptase to make DNA copies of mature mRNA isolated from eukaryotic cells. These copies, known as complementary DNA (cDNA), are then incorporated into recombinant DNA vectors. Learn more about the formation of cDNA by clicking on the figure in CengageNOW.

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The polymerase chain reaction (PCR) is a widely used, automated, in vitro technique in which researchers use specific primers to target a particular DNA sequence and then clone it using a heatresistant DNA polymerase. Using PCR, scientists amplify and analyze tiny DNA samples taken from various sites, from crime scenes to archaeological remains. Watch the amplification of DNA by PCR by clicking on the figure in CengageNOW.

15.2 (page 331) 5 Distinguish among DNA, RNA, and protein blotting. A Southern blot detects DNA fragments by separating them using gel electrophoresis and then transferring them to a nitrocellulose or nylon membrane. A probe is then hybridized by complementary base pairing to the DNA bound to the membrane, and the band or bands of DNA are identified by autoradiography or chemical luminescence. ■ When RNA molecules that are separated by electrophoresis are transferred to a membrane, the result is a Northern blot. ■ A Western blot consists of proteins or polypeptides previously separated by gel electrophoresis. 6 Describe the chain termination method of DNA sequencing. ■ DNA sequencing yields information about the structure of a gene and the probable amino acid sequence of its encoded proteins. ■ Automated DNA sequencing is based on the chain termination method, which uses dideoxynucleotides, each tagged with a different-colored fluorescent dye, to terminate elongation during DNA replication. Gel electrophoresis separates the resulting fragments, and a laser identifies the nucleotide sequence. ■

Learn more about automated DNA sequencing by clicking on the figure in CengageNOW.

15.3 (page 335)

causing genes or the proteins they code for, which can then be targeted by therapeutic drugs. 9 Define pharmacogenetics and proteomics. ■ Pharmacogenetics, the new science of gene-based medicine, takes into account the subtle genetic differences among individuals and customizes drugs to match a patient’s genetic makeup. ■ The study of all the proteins encoded by the human genome and produced in a person’s cells and tissues is proteomics. Scientists want to identify all the proteins made by a given kind of cell, but the process is much more complicated than sequencing the human genome.

15.4 (page 339) 10 Describe at least one important application of recombinant DNA technology in each of the following fields: medicine, DNA fingerprinting, and transgenic organisms. ■ Genetically altered bacteria produce many important human protein products, including insulin, growth hormone, tissue plasminogen activator (TPA), tissue growth factor-beta (TGF-b), clotting factor VIII, and Dornase Alpha (DNase). ■ DNA fingerprinting is the analysis of an individual’s DNA. It is based on a variety of short tandem repeats (STRs), molecular markers that are highly polymorphic within the human population. DNA fingerprinting has applications in areas such as law enforcement, issues of disputed parentage, and tracking tainted foods.

7 Describe three important areas of research in genomics. Genomics is the emerging field of biology that studies the entire DNA sequence of an organism’s genome. Structural genomics is concerned with mapping and sequencing genomes. Functional genomics deals with the functions of genes and nongene sequences in genomes. Comparative genomics involves comparing the genomes of different species to further our understanding of evolutionary relationships. 8 Explain what a DNA microarray does and give an example of its research and medical potential. ■ Many diagnostic tests involve DNA microarrays, in which hundreds of different DNA molecules are placed on a glass slide or chip. DNA microarrays enable researchers to compare the activities of thousands of genes in normal and diseased cells from tissue samples. Because cancer and other diseases exhibit altered patterns of gene expression, DNA microarrays have the potential to identify disease-

Solve a murder case with DNA fingerprinting by clicking on the figure in CengageNOW.

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Transgenic organisms have foreign DNA incorporated into their genetic material. Transgenic livestock produce foreign proteins in their milk. Transgenic plants have great potential in agriculture.

15.5 (page 343) 11 Describe at least two safety issues associated with recombinant DNA technology and explain how these issues are being addressed. ■ Some people are concerned about the safety of molecularly modified organisms. To address these concerns, scientists carry out recombinant DNA technology under specific safety guidelines. ■ The introduction of transgenic plants and animals into the natural environment, where they may spread in an uncontrolled manner, is an ongoing concern.

T E S T YO U R U N D E R S TA N D I N G 1. A plasmid (a) can be used as a DNA vector (b) is a type of bacteriophage (c) is a type of cDNA (d) is a retrovirus (e) b and c

5. The PCR technique uses (a) heat-resistant DNA polymerase (b) reverse transcriptase (c) DNA ligase (d) restriction enzymes (e) b and c

2. DNA molecules with complementary sticky ends associate by (a) covalent bonds (b) hydrogen bonds (c) ionic bonds (d) disulfide bonds (e) phosphodiester linkages

6. A cDNA clone contains (a) introns (b) exons (c) anticodons (d) a and b (e) b and c

3. Human DNA and a particular plasmid both have sites that are cut by the restriction enzymes HindIII and EcoRI. To make recombinant DNA, the scientist should (a) cut the plasmid with EcoRI and the human DNA with HindIII (b) use EcoRI to cut both the plasmid and the human DNA (c) use HindIII to cut both the plasmid and the human DNA (d) a or b (e) b or c

7. The dideoxynucleotides ddATP, ddTTP, ddGTP, and ddCTP are important in DNA sequencing because they (a) cause premature termination of a growing DNA strand (b) are used as primers (c) cause the DNA fragments that contain them to migrate more slowly through a sequencing gel (d) are not affected by high temperatures (e) have more energy than deoxynucleotides

4. Which technique rapidly replicates specific DNA fragments without cloning in cells? (a) gel electrophoresis (b) cDNA libraries (c) DNA probe (d) restriction fragment length polymorphism (e) polymerase chain reaction

8. In the Southern blot technique, _____ is/are transferred from a gel to a nitrocellulose or nylon membrane. (a) protein (b) RNA (c) DNA (d) bacterial colonies (e) reverse transcriptase

9. Gel electrophoresis separates nucleic acids on the basis of differences in (a) length (molecular weight) (b) charge (c) nucleotide sequence (d) relative proportions of adenine and guanine (e) relative proportions of thymine and cytosine 10. A genomic DNA library (a) represents all the DNA in a specific chromosome (b) is made using reverse transcriptase (c) is stored in a collection of recombinant bacteria (d) is a DNA copy of mature mRNAs (e) allows researchers to amplify a tiny sample of DNA

11. Tissue growth factor-beta (a) is a DNA probe for recombinant plasmids (b) is a product of DNA technology used in tissue engineering (c) is necessary to make a cDNA library (d) cannot be synthesized without a heat-resistant DNA polymerase (e) is isolated by the Southern blot technique 12. These highly polymorphic molecular markers are useful in DNA fingerprinting: (a) cosmid cloning vectors (b) cloned DNA sequences (c) palindromic DNA sequences (d) short tandem repeats (e) complementary DNAs

CRITICAL THINKING 1. What are some of the problems that might arise if you were trying to produce a eukaryotic protein in a bacterium? How might using transgenic plants or animals help solve some of these problems? 2. Would molecular modification be possible if we did not know a great deal about the genetics of bacteria? Explain. 3. How is proteomics related to functional genomics?

Leonard Lessin/Peter Arnold, Inc.

4. ANALYZING DATA. RFLP analysis was performed on blood from a crime scene as well as from two suspects. The crime scene DNA is marked E(vs). Which suspect is probably guilty? Which suspect is definitely innocent?

5. EVOLUTION LINK. DNA technology, such as the production of transgenic animals, is possible only because widely different organisms have essentially identical genetic systems (DNA ¡ RNA ¡ protein). What is the evolutionary significance of the universality of genetic systems in organisms as diverse as bacteria and pigs? 6. SCIENCE, TECHNOLOGY, AND SOCIETY. What are some ways that recent DNA technologies might affect you personally? Additional questions are available in CengageNOW at www.cengage.com/ login.

Human Genetics and the Human Genome

© Regents of the University of California 2005/Dr. Uli Weier/Photo Researchers, Inc.

16

KEY CONCEPTS 16.1 Pedigree analysis and karyotype analysis are traditional methods to study human genetics; molecular aspects of the human genome, including genome-wide association scans and comparative genomics, are increasingly used today. 16.2 Chromosome abnormalities such as aneuploidies—

Rapid chromosome mapping. This fluorescent LM of human chromosome 10 (left) has different-colored fluorescent probes attached to specific DNA sequences. (Loci and key are at right.) This technique allows researchers to efficiently determine certain chromosome rearrangements and abnormalities.

T

he principles of genetics apply to all organisms, including humans. However, important differences separate genetic research on humans

and genetic research on other organisms. To study inheritance in other species, geneticists may conduct controlled matings between individuals and raise the offspring under carefully controlled conditions. Of course, setting up experimental matings in humans is unethical and illegal. An important difference between genetic research in humans and

the presence or absence of a single chromosome—can cause serious medical conditions.

other organisms is that organisms used in genetic research generally

16.3 Mutations in single genes can produce genetic

ductive periods. In contrast, most human families are small, and 20 to

disorders.

30 years or more elapse between generations.

16.4 Researchers are developing gene therapy methods that could potentially correct certain genetic disorders.

16.5 Genetic testing and counseling help individuals make reproductive decisions.

16.6 Certain advances in human genetics raise ethical questions.

produce many offspring and have short generation times between repro-

Despite the inherent difficulties of studying inheritance in humans, human genetics, the science of inherited variation in humans, is progressing rapidly (see figure). Researchers conduct population studies of large extended families. Also, the medical attention given to human genetic diseases has expanded our knowledge of human genetics. Genetic studies of other organisms have provided invaluable insights: many

human inheritance questions have been explained using model organisms, such as bacteria, yeasts, worms, fruit flies, and mice. More recently, the human genome, which represents the totality of genetic information in human cells, has been mapped and sequenced. In DNA sequencing, researchers identify the order of nucleotides in DNA to understand the genetic basis of human similarities and differences. In this chapter we first examine the methods of human genetics, including new research directions that are developing as a result of the Human Genome Project. Then we discuss a variety of human genetic disorders. We explore the use of gene therapy for some of these disorders, as well as the application of genetic testing, screening, and counseling for families at risk. The chapter concludes with a discussion of ethical issues related to human genetics.

16.1 STUDYING HUMAN GENETICS ■ ■ LEARNING OBJECTIVES 1 2 3

Distinguish between karyotyping and pedigree analysis. Discuss the implications of the Human Genome Project. Discuss the mouse model for studying cystic fibrosis.

Human geneticists use a variety of methods that enable them to help identify genetic defects and to make inferences about a trait’s mode of inheritance. We consider three of these methods—the identification of chromosomes by karyotyping, the analysis of family inheritance patterns using pedigrees, and DNA sequencing and mapping of genes by genome projects. Investigators often study human inheritance most effectively by combining these and other approaches.

Human chromosomes are studied by karyotyping Like Mendel and his pea plants, researchers working with simpler organisms have discovered many of the basic principles of genetics. In such organisms, it is often possible to relate genetic data to the number and structure of specific chromosomes. Some organisms used in genetics, such as the fruit fly Drosophila melanogaster, have very few chromosomes; the fruit fly has only four pairs. In Drosophila larval salivary glands, the chromosomes are large enough that their structural details are readily evident. (Note the detail shown in the light micrograph of Drosophila chromosomes in Figure 2-3.) This organism, therefore, has provided unique opportunities for correlating certain inherited phenotypic changes with alterations in chromosome structure. The normal number of chromosomes for the human species is 46: 44 autosomes (22 pairs) and 2 sex chromosomes (1 pair). A karyotype (from Greek, meaning “nucleus”) is an individual’s chromosome composition, both chromosome number and large structural defects in a chromosome. Until the mid-1950s, when biologists adopted modern methods of karyotyping, the accepted

number of chromosomes for the human species was 48, based on a study of human chromosomes published in 1923. The reason researchers counted 48 human chromosomes was the difficulty in separating the chromosomes so they could be accurately counted. In 1952, University of Texas cell biologist T. C. Hsu treated cells with a hypotonic salt solution by mistake; this caused the cells to swell and the chromosomes to spread apart, thereby making them easier to count. Other techniques were also developed, and in 1956 researchers Joe Hin Tjio and Albert Levan, working in Sweden, reported that humans have 46 chromosomes, not 48. Other researchers subsequently verified this report. The story of the human chromosome number is a valuable example of the “selfcorrecting” nature of science (although such corrections may take time). The re-evaluation of established facts and ideas, often by using improved techniques or new methods, is an essential part of the scientific process. Human chromosomes are visible only in dividing cells (see Chapter 10), and it is difficult to obtain dividing cells directly from the human body. Researchers typically use blood because white blood cells can be induced to divide in a culture medium by treating them with certain chemicals. Other sources of dividing cells include skin and, for prenatal chromosome studies, chorionic villi or fetal cells shed into the amniotic fluid (discussed later in the chapter). In karyotyping, biologists culture dividing human cells and then treat them with the drug colchicine, which arrests the cells at mitotic metaphase or late prophase, when the chromosomes are most highly condensed. Next, the researchers put the cells into a hypotonic solution; the cells swell, and the chromosomes spread out so they are easily observed. The investigators then flatten the cells on microscope slides and stain the chromosomes to reveal the patterns of bands, which are unique for each homologous pair. After the microscopic image has been scanned into a computer, the homologous pairs are electronically matched and placed together (FIG. 16-1). By convention, geneticists identify chromosomes by length; position of the centromere; banding patterns, which are produced by staining chromosomes with dyes that produce dark and light crossbands of varying widths; and other features such as satellites, tiny knobs of chromosome material at the tips of certain chromosomes. With the exception of the sex chromosomes, all chromosomes but chromosome 21, which is smaller than chromosome 22, are numbered and lined up in order of size. The largest human chromosome (chromosome 1) is about five times as long as the smallest one (chromosome 21), but there are only slight size differences among some of the intermediate-sized chromosomes. The X and Y chromosomes of a normal male are homologous only at their tips; normal females have two X chromosomes and no Y chromosome. Differences from the normal karyotype—that is, deviations in chromosome number or structure—are associated with certain disorders such as Down syndrome (discussed later in the chapter). Another way to distinguish chromosomes in a karyotype is by fluorescent in situ hybridization (FISH). A geneticist tags a DNA strand complementary to DNA in a specific chromosome with a fluorescent dye. The DNA in the chromosome is denatured—that is, the two strands are separated—so the tagged

strand can bind to it. A different dye is used for each chromosome, “painting” each with a different color (see karyotype in Fig. 16-1). A chromosome that is multicolored (not shown) indicates breakage and fusion of chromosomes, an abnormality associated with certain genetic diseases and many types of cancer. RESEARCH METHOD Researchers use karyotypes to screen individuals for chromosome abnormalities.

How Is It Done?

Karyotypes are commonly prepared from cultures of white blood cells. Researchers place prepared cells on a slide, and the chromosomes spill out of the ruptured nuclei. The light microscope under which the slide is viewed is connected to a computer with image analysis software, which helps the biologist match homologous chromosomes and organize them by size. Before computers were used, researchers cut and pasted chromosomes from photographs.

SIU/Peter Arnold, Inc.

Why Is It Used?

© Dr. Thomas Liehr, courtesy of Biology Image Library

These chromosomes have been “painted” by different-colored fluorescent dyes that hybridize to specific pairs of homologous chromosomes. Painting helps the researcher identify chromosomes. A normal human male karyotype is shown.

FIGURE 16-1 Karyotyping

Family pedigrees help identify certain inherited conditions Early studies of human genetics usually dealt with readily identified pairs of contrasting traits and their distribution among members of a family. A “family tree” that shows inheritance patterns, the transmission of genetic traits within a family over several generations, is known as a pedigree. Pedigree analysis remains widely used, even in today’s world of powerful molecular genetic techniques, because it helps molecular geneticists determine the exact interrelationships of the DNA molecules they analyze from related individuals. Pedigree analysis is also an important tool of genetic counselors and clinicians. However, because human families tend to be small and information about certain family members, particularly deceased relatives, may not be available, pedigree analysis has limitations. Pedigrees are produced using standardized symbols. Examine FIGURE 16-2, which shows a pedigree for albinism, a lack of the pigment melanin in the skin, hair, and eyes. Each horizontal row represents a separate generation, with the earliest generation (Roman numeral I) at the top and the most recent generation at the bottom. Within a given generation, the individuals are usually numbered consecutively, from left to right, using Arabic numerals. A horizontal line connects two parents, and a vertical line drops from the parents to their children. For example, individuals II-3 and II-4 are parents of four offspring (III-1, III-2, III-3, and III-4). Note that individuals in a given generation can be genetically unrelated. For example, II-1, II-2, and II-3 are unrelated to II-4 and II-5. Within a group of siblings, the oldest is on the left, and the youngest is on the right. The allele for albinism cannot be dominant; if it were, at least one of female III-2’s parents would have been an albino. This pedigree is explained if albinism is inherited as an autosomal (not carried on a sex chromosome) recessive allele. In such cases, two phenotypically normal parents could produce an albino offspring because they are heterozygotes and could each transmit a recessive allele. Studying pedigrees enables human geneticists to predict how phenotypic traits that are governed by the genotype at a single locus are inherited. About 10,000 traits have been described in humans. Pedigree analysis most often identifies three modes of single-locus inheritance: autosomal dominant, autosomal recessive, and X-linked recessive. We define and discuss examples of these inheritance modes later in the chapter.

The Human Genome Project sequenced the DNA on all human chromosomes Scientists working on the Human Genome Project sequenced the DNA in the entire nuclear human genome—about 2.9 billion base pairs. (The human mitochondrial genome was sequenced in 1981.) This international undertaking, based on the DNA from 6 to 10 anonymous individuals, was essentially completed in 2001 by hundreds of researchers working in two independent teams, the government-funded International Human Genome Sequencing Consortium and the privately funded company Celera Genomics. The completion of the Human Genome Project in 2003 was a significant milestone in genetics (TABLE 16-1).

©Mark Eveleigh/Alamy

TABLE 16-1

(a) An albino Cuna Indian girl plays with her friends. Albinism is common among the Cuna people living on the San Blas Islands of Panama.

Selected Milestones in Genetics

Year

Scientific Advance

1866

Mendel proposed existence of hereditary factors now known as genes

1871

Nucleic acids discovered

1879

Chromosomes identified and observed during cell division

1929

Chemical nature of nucleotides characterized

1953

Double-helix structure of DNA determined

1960s

Genetic code explained (how proteins are made from DNA)

1972

First recombinant DNA molecule produced

1977

DNA sequencing began

1986

DNA sequencing automated

1995

Sequencing of first prokaryotic genome (bacterium Haemophilus influenzae) completed

1996

Sequencing of first eukaryotic genome (yeast Saccharomyces cerevisiae) completed

1998

Sequencing of first multicellular eukaryotic genome (nematode worm Caenorhabditis elegans) completed

2001

Draft sequence of entire human genome published; discovered that 95% of genome is noncoding

2003

Completion of DNA sequencing of human genome

I 1

2

3

4

II 1

2

3

4

5

III 1

2

3

4

Key: Normal female

Mating

Normal male Albino female Albino male

Siblings produced by mating

(b) By studying family histories, a researcher can determine the genetic mechanism of an inherited trait. In this example, III-2 represents an albino girl with two phenotypically normal parents, II-3 and II-4.

FIGURE 16-2 Animated A pedigree for albinism

Scientists hope to eventually identify where all the genes are located in the sequenced DNA. In 2001, the National Human Genome Research Institute (NHGRI) estimated there were 35,000 to 45,000 protein-coding genes in the human genome, but by the end of the decade the estimate had been lowered to 20,000 to 25,000.

Gene identification represents a formidable challenge. How do you identify a gene in a DNA sequence if you know nothing about it? The human genome is extremely complex. It includes the DNA content of both the nucleus and the mitochondria. However, nuclear DNA accounts for almost all the genetic information in the human genome. Like genomes of other eukaryotic organisms, some (in humans, only about 2%) of the human genome specifies the synthesis of polypeptides, whereas some other segments code only for RNA products. Most of the human genome consists of noncoding regulatory elements, repetitive sequences (multiple copies) of DNA, and gene segments whose functions remain unknown. Sometimes genes overlap one another, and in other cases a single gene codes for multiple proteins. Relating specific genes to the RNAs and proteins they code for and determining the roles these RNAs and proteins play in the body are some of the avenues of human genetic research that scientists are currently pursuing. In human chromosome 22, for example, computers scanned DNA sequences for identifying markers such as promoter elements that are typically associated with genes. Using this approach, researchers identified 545 genes. Many of these are “potential” genes in that their messenger RNAs (mRNAs) and protein products have not yet been isolated. Other genes were identified because they code for proteins similar to those previously identified in humans or other organisms. Despite the difficulties, scientists are making

progress, and mapping studies will help scientists understand the physical and functional relationships among genes and groups of genes as revealed by their order on the chromosomes. Now that the human genome has been sequenced, researchers will be busy for many decades analyzing the growing body of human molecular data. In addition to identifying genes, scientists want to understand what each gene’s role is, how each gene interacts with other genes, and how the expression of each gene is regulated in different tissues. Eventually, all the thousands of proteins produced in human cells will be identified, their 3-D structures determined, and their properties and functions evaluated. Genome-wide association scans demonstrate the genetic complexity of major human diseases Researchers have begun to study sequence variations within the human genome to elucidate differences that might be related to susceptibility to illness or disease. The potential medical applications of the Human Genome Project are extremely promising. Genes on human chromosome 22, for example, are associated with at least 27 diseases known to have a genetic component. The causative genes of many of these disorders have not yet been identified. For example, a gene involved in schizophrenia is strongly linked to human chromosome 22, but scientists do not know its exact location or function. Genome-wide association (GWA) scans compare the genomes of individuals with a particular disease to healthy individuals without that disease. In 2007, for example, researchers scanned the genomes of more than 16,000 British people to compare genetic patterns in healthy people with those in people with one of seven common diseases: bipolar disorder, coronary artery disease, Crohn’s disease, hypertension, rheumatoid arthritis, type 1 diabetes, and type 2 diabetes. These disease phenotypes exhibit complex inheritance patterns, with many different genes contributing to susceptibility. In this particular study, scientists found 24 variations associated with an increased risk of one or more of these diseases. The functions of most of these GWA variations have not yet been determined, but knowing their locations on particular chromosomes will make it easier for researchers to study the genetics of many common human diseases.

Comparative genomics has revealed DNA identical in both mouse and human genomes About 500 DNA segments longer than 200 base pairs are known to be 100% conserved (that is, identical) between the mouse and human genomes. This remarkable degree of conservation has interesting evolutionary implications because it means that these segments have not mutated during the past 75 million years that mice and humans have shared a common ancestor. During this time, other DNA sections underwent considerable mutation and selection, allowing mice and humans to diverge to their present states. Although the functions of these unchanged elements are not yet determined, they clearly have a vital role. (If they were not essential in their present form, they would have undergone mutation and selection.) Many highly conserved segments appear to contain nonprotein-coding elements that may regulate the expression of other genes.

Researchers use mouse models to study human genetic diseases Many questions relating to human genetic diseases are difficult to answer because of the ethical issues involved in using humans as test subjects. However, research on any disease is greatly facilitated if an animal model is used for experimentation. A good example is cystic fibrosis, a genetic disease caused by a single gene mutation inherited as a recessive allele. In 1994, researchers used gene targeting to produce strains of mice that were either homozygous or heterozygous for cystic fibrosis. The allele that causes cystic fibrosis is a mutant form of a locus involved in controlling the body’s water and electrolyte balance. Geneticists have cloned this gene and found that it codes for a protein, the CFTR protein, that serves as a chloride ion channel in the plasma membrane. (CFTR stands for cystic fibrosis transmembrane conductance regulator.) This ion channel transports chloride ions out of the cells lining the digestive tract and the respiratory system. When the chloride ions leave the cells, water follows by osmosis. Thus, the normal secretions of these cells are relatively watery. Because the cells of individuals with cystic fibrosis lack normal chloride ion channels, the individuals’ secretions have a low water content and their sweat is very salty. Cells of heterozygous individuals have only half the usual number of functional CFTR ion channels, but these are enough to maintain the normal fluidity of their secretions. Some researchers are now focusing on understanding the way in which the CFTR channel is activated or inactivated in mice. They hope to use this information to design drugs that enhance chloride transport through the CFTR channels. Such drugs have the potential to treat cystic fibrosis in humans by activating the mutant channels.

Review ■ ■ ■

■

What kinds of information can a human karyotype provide? What is pedigree analysis? What are two possible benefits scientists hope to obtain by further study of the human genome? How does using a mouse model for a genetic disease overcome some of the difficulties in studying human inheritance?

16.2 ABNORMALITIES IN CHROMOSOME NUMBER AND STRUCTURE ■ ■ LEARNING OBJECTIVES 4 Explain how nondisjunction in meiosis is responsible for chromosome

5

abnormalities such as Down syndrome, Klinefelter syndrome, and Turner syndrome. Distinguish among the following structural abnormalities in chromosomes: translocations, deletions, and fragile sites.

Polyploidy, the presence of multiple sets of chromosomes, is common in plants but rare in animals. It may arise from the failure of chromosomes to separate during meiosis or from the fertilization

of an egg by more than one sperm. When it occurs in all the cells of the body, polyploidy is lethal in humans and many other animals. For example, triploidy (3n) is sometimes found in human embryos that have been spontaneously aborted in early pregnancy. Abnormalities caused by the presence of a single extra chromosome or the absence of a chromosome—called aneuploidies— are more common than polyploidy. Disomy is the normal state: two of each kind of chromosome in a cell or individual. In trisomy, a cell or individual has two copies of each chromosome except for one, which has three copies; a trisomic chromosome number is designated 2n + 1. In monosomy, an individual lacks one member of a pair of chromosomes; a monosomic chromosome number is designated 2n − 1. TABLE 16-2 summarizes some disorders that aneuploidies produce. Aneuploidies generally arise as a result of an abnormal meiotic (or, rarely, mitotic) division in which chromosomes fail to separate at anaphase. This phenomenon, called nondisjunction, can occur with the autosomes or with the sex chromosomes. In meiosis, chromosome nondisjunction may occur during the first or second meiotic division (or both). For example, two X chromosomes that fail to separate at either the first or the second meiotic division may both enter the egg nucleus. Alternatively, the two joined X chromosomes may go into a polar body, leaving the egg with no X chromosome. (Recall from Chapter 10 that a polar body is a nonfunctional haploid cell produced during oogenesis; also see Fig. 50-11.) Nondisjunction of the XY pair during the first meiotic division in the male may lead to the formation of a sperm with both X and Y chromosomes or a sperm with neither an X nor a Y chromosome (FIG. 16-3). Similarly, nondisjunction at the second meiotic division can produce sperm with two Xs or two Ys. When an abnormal gamete unites with a normal one, the resulting zygote has a chromosome abnormality that will be present in every cell of the body. Meiotic nondisjunction results in an abnormal chromosome number at the zygote stage of development, so all cells in the in-

TABLE 16-2

dividual have an abnormal chromosome number. In contrast, nondisjunction during a mitotic division occurs sometime later in development and leads to the establishment of a clone of abnormal cells in an otherwise normal individual. Such a mixture of cells with different chromosome numbers may or may not affect somatic (body) or germ line (reproductive) tissues. Different numbers of chromosomes are common in many cancer cells, particularly those in solid tumors. Whether aneuploidy is a cause or a consequence of cancer, however, is not yet clear. Recognizable chromosome abnormalities are seen in less than 1% of all live births, but substantial evidence suggests the rate at conception is much higher. At least 17% of pregnancies recognized at 8 weeks will end in spontaneous abortion (miscarriage). Approximately half of these spontaneously aborted embryos have major chromosome abnormalities, including autosomal trisomies (such as trisomy 21), triploidy, tetraploidy, and Turner syndrome (X0), in which the 0 refers to the absence of a second sex chromosome. Autosomal monosomies are exceedingly rare, possibly because they induce a spontaneous abortion very early in the pregnancy, before a woman is even aware she is pregnant. Some investigators give surprisingly high estimates (50% or more) for the loss rate of very early embryos. Chromosome abnormalities probably induce many of these spontaneous abortions.

Down syndrome is usually caused by trisomy 21 Down syndrome is one of the most common chromosome abnormalities in humans. (The term syndrome refers to a set of symptoms that usually occur together in a particular disorder.) It was named after J. Langdon Down, the British physician who in 1866 first described the condition. Affected individuals have abnormalities of the face, eyelids, tongue, hands, and other parts of the body and are mentally retarded (FIG. 16-4a). They are also unusually susceptible to certain diseases, such as leukemia and Alzheimer’s disease.

Chromosome Abnormalities: Disorders Produced by Aneuploidies

Karyotype

Common Name

Clinical Description

Trisomy 13

Patau syndrome

Multiple defects, with death typically by age 3 months

Trisomy 18

Edwards syndrome

Ear deformities, heart defects, spasticity, and other damage; death typically by age 1 year, but some survive much longer

Trisomy 21

Down syndrome

Overall frequency is about 1 in 800 live births. Most conceptions involving true trisomy occur in older (age 35+) mothers, but translocation resulting in the equivalent of trisomy is not age-related. Trisomy 21 is characterized by a fold of skin above the eye, varying degrees of mental retardation, short stature, protruding furrowed tongue, transverse palmar crease, cardiac deformities, and increased risk of leukemia and Alzheimer’s disease.

X0

Turner syndrome

Short stature, webbed neck, sometimes slight mental retardation; ovaries degenerate in late embryonic life, leading to rudimentary sexual characteristics; gender is female; no Barr bodies

XXY

Klinefelter syndrome

Male with slowly degenerating testes, enlarged breasts; one Barr body per cell

XYY

XYY karotype

Many males have no unusual symptoms; others are unusually tall, with heavy acne, and some tendency to mild mental retardation

XXX

Triplo-X

Despite three X chromosomes, usually fertile females with normal intelligence; two Barr bodies per cell

KEY POINT

Aneuploidy may occur by meiotic nondisjunction, the abnormal segregation of chromosomes during meiosis.

Nondisjunction in first meiotic division

XY

XY X Y

(a) First meiotic division nondisjunction results in two XY sperm and two sperm with neither an X nor a Y. Nondisjunction of X in second meiotic division Normal first meiotic division

Nondisjunction of Y in second meiotic division XX

X

X X Y Y

Y

YY

(b) Second meiotic division nondisjunction of the X chromosome results in one sperm with two X chromosomes, two with one Y each, and one with no sex chromosomes. Nondisjunction of the Y chromosome results in one sperm with two Y chromosomes, two with one X each, and one with no sex chromosome (box on right).

FIGURE 16-3 Meiotic nondisjunction In these examples of nondisjunction of the sex chromosomes in the human male, only the X (purple) and Y (blue) chromosomes are shown. (The positions of the metaphase chromosomes have been modified to save space.)

Genetic studies have revealed that most people with Down syndrome have 47 chromosomes because of autosomal trisomy: this condition is known as trisomy 21 (FIG. 16-4b). Nondisjunction during meiosis is responsible for the presence of the extra chromosome. Although no genetic information is missing in these individuals, the extra copies of chromosome 21 genes bring about some type of genetic imbalance that causes abnormal physical and mental development. Down syndrome is quite variable in expression, with some individuals far more severely affected than others. Researchers are using DNA technologies to pinpoint genes on chromosome 21 that affect mental development, as well as possible oncogenes (cancer-causing genes) and genes that may be

involved in Alzheimer’s disease. (Like many human conditions, cancer and Alzheimer’s disease have both genetic and environmental components.) Down syndrome occurs in all ethnic groups in about 1 out of 800 live births. Its incidence increases markedly with increasing maternal age. The occurrence of Down syndrome is not affected by the father’s age (although other disorders are, including schizophrenia and achondroplasia, the most common form of dwarfism). Down syndrome is 68 times as likely in the offspring of mothers of age 45 than in the offspring of mothers age 20. However, most babies with Down syndrome in the United States are born to mothers younger than 35, in part because these greatly outnumber older mothers, and

CNRI/Science Photo Library/Photo Researchers, Inc.

Richard Hutchings/Photo Researchers, Inc.

(a) This boy with Down syndrome is working on a science experiment in his kindergarten class. Some individuals with Down syndrome learn to read and write.

(b) Note the presence of an extra chromosome 21 in this colorized karyotype of a female with Down syndrome.

FIGURE 16-4 Down syndrome

in part because about 90% of older women who undergo prenatal testing terminate the pregnancy if Down syndrome is diagnosed. The relationship between increased incidence in Down syndrome and maternal age has been studied for decades, but there is no explanation. Scientists have proposed several hypotheses to explain the maternal age effect, but none are supported unequivocally. One explanation is that older women have held eggs in suspended meiosis too long, leading to a deterioration of the meiotic spindle apparatus. (A woman is born with all the oocytes she will ever have; these oocytes remain in prophase I of meiosis until ovulation.) Another possibility is that an aging uterus is less likely to reject an abnormal fetus.

Most sex chromosome aneuploidies are less severe than autosomal aneuploidies Sex chromosome aneuploidies are tolerated relatively well (see Table 16-2) This is true, at least in part because of the mechanism of dosage compensation: mammalian cells compensate for extra X chromosome material by rendering all but one X chromosome inactive. The inactive X is seen as a Barr body, a region of darkly stained, condensed chromatin next to the nuclear envelope of an interphase nucleus (see Fig. 11-16). Investigators have used the presence of the Barr body in the cells of normal females as an initial screen to determine whether an individual is genetically female or male. However, as you will see shortly in the context of sex chromosome aneuploidies, the Barr body test has limitations. Individuals with Klinefelter syndrome are males with 47 chromosomes, including two Xs and one Y. They have small testes,

produce few or no sperm, and are therefore sterile. The hypothesis that the Y chromosome is the major determinant of the male phenotype has been substantiated by the fact that at least one gene on the Y chromosome acts as a genetic switch, directing male development. Males with Klinefelter syndrome tend to be unusually tall and have female-like breast development. About half show some mental retardation, but many live relatively normal lives. However, each of their cells has one Barr body. On the basis of such a test, they would be erroneously classified as females. About 1 in 1000 live-born males has Klinefelter syndrome. The sex chromosome composition for Turner syndrome, in which an individual has only one sex chromosome, an X chromosome, is designated X0. Because they lack the male-determining effect of the Y chromosome, individuals with Turner syndrome develop as females. However, both their internal and their external genital structures are underdeveloped, and they are sterile. Apparently, a second X chromosome is necessary for normal development of ovaries in a female embryo. Examination of the cells of these individuals reveals no Barr bodies because there is no extra X chromosome to be inactivated. Using the standards of the Barr body test, such an individual would be classified erroneously as a male. About 1 in 2500 live-born females has Turner syndrome. People with an X chromosome plus two Y chromosomes are phenotypically males, and they are fertile. Other characteristics of these individuals (tall, often with severe acne) hardly qualify as a syndrome; hence the designation XYY karyotype. Some years ago, several widely publicized studies suggested that males with this condition are more likely to display criminal tendencies and thus to be imprisoned. However, these studies were flawed because

they were based on small numbers of XYY males, without adequate or well-matched control studies of XY males. The prevailing opinion in medical genetics today is that there are many undiagnosed XYY males in the general population who do not have overly aggressive or criminal behaviors and are unlikely to be incarcerated.

Translocation is the attachment of part of one chromosome to another The consequences of translocations vary considerably. They include deletions, in which some genes are missing, and duplications, in which extra copies of certain genes are present. In about 4% of individuals with Down syndrome, only 46 chromosomes are present, but one is abnormal. The large arm of chromosome 21 has been translocated to the large arm of another chromosome, usually chromosome 14. Individuals with translocation Down syndrome have one chromosome 14, one combined 14/21 chromosome, and two normal copies of chromosome 21. All or part of the genetic material from chromosome 21 is thus present in triplicate. When studying the karyotypes of the parents in such cases, geneticists usually find that either the mother or the father has only 45 chromosomes, although she or he is generally phenotypically normal. The parent with 45 chromosomes has one chromosome 14, one combined 14/21 chromosome, and one chromosome 21; although the karyotype is abnormal, there is no extra genetic material. In contrast to trisomy 21, translocation Down syndrome can run in families, and its incidence is not related to maternal age.

Abnormalities in chromosome structure cause certain disorders Chromosome abnormalities are caused not only by changes in chromosome number but also by distinct changes in the structure of one or more chromosomes. The breakage and rejoining of chromosome parts result in four structural changes within or between chromosomes: duplications, inversions, deletions, and translocations (FIG. 16-5). Breaks in chromosomes are the result of errors in replication or recombination. In a duplication, a segment of the chromosome is repeated one or more times; these repeats often appear in tandem with one another. The orientation of a chromosome segment is reversed in an inversion. In a deletion, breakage causes loss of part of a chromosome, along with the genes on that segment. A deletion can occur at the end of a chromosome or on an internal part of the chromosome. In some cases of translocation, a chromosome fragment breaks off and attaches to a nonhomologous chromosome. In a reciprocal translocation, two nonhomologous chromosomes exchange segments. Here we consider three simple examples of chromosomal structural changes that result in abnormal phenotypes: translocations, deletions, and fragile sites, which are chromosome sites that are susceptible to breakage.

A deletion is the loss of part of a chromosome Sometimes chromosomes break and fail to rejoin. Such breaks result in deletions of as little as a few base pairs to as much as an entire chromosome arm. As you might expect, large deletions are generally lethal, whereas small deletions may have no effect or may cause recognizable human disorders. One deletion disorder (1 in 50,000 live births) is cri du chat syndrome, in which part of the short arm of chromosome 5 is

Lost segment

A

A

A

A

B

B

A

A A A B

A

(a) A duplication is a repeated segment of a chromosome. In this example, segment A is repeated.

A

(b) An inversion is a chromosome segment with a reversed orientation. An inversion does not change the amount of genetic material in the chromosome, only its arrangement.

(c) A deletion is the loss of a chromosome segment. A deletion can occur at the tip (shown) or within the chromosome.

FIGURE 16-5 Animated Common abnormalities of chromosome structure

B A

(d) A reciprocal translocation occurs when two nonhomologous chromosomes exchange segments.

1 μm

Fragile sites are weak points at specific locations in chromatids A fragile site is a place where part of a chromatid appears to be attached to the rest of the chromosome by a thin thread of DNA. Fragile sites occur at corresponding locations on both chromatids of a chromosome. They have been identified on the X chromosome as well as on certain autosomes. The location of a fragile site is exactly the same in all of an individual’s cells, as well as in cells of other family members. Scientists report growing evidence that cancer cells may have breaks at these fragile sites. Whether cancer destabilizes the fragile sites, leading to breakage, or the fragile sites themselves contain genes that contribute to cancer is unknown at this time. In fragile X syndrome, also known as Martin–Bell syndrome, a fragile site occurs near the tip of the X chromosome, where the fragile X gene contains a nucleotide triplet CGG that repeats from 200 to more than 1000 times (FIG. 16-6). In a normal chromosome, CGG repeats up to 50 times. Fragile X syndrome is the most common cause of inherited mental retardation. The effects of fragile X syndrome, which are more pronounced in males than in females, range from mild learning and attention deficit disorders to severe mental retardation and hyperactivity. According to the National Fragile X Foundation, about 80% of boys and 35% of girls with fragile X syndrome are at least mildly mentally retarded. Females with fragile X syndrome are usually heterozygous (because their other X chromosome is normal) and are therefore more likely to have normal intelligence. The discovery of the fragile X gene in 1991 and the development of the first fragile X mouse model in 1994 have provided researchers with ways to develop and test potential treatments, including gene therapy. At the microscopic level, the nerve cells of individuals with fragile X syndrome have malformed dendrites (the part of the nerve cell that receives nerve impulses from other nerve cells). At the molecular level, the triplet repeats associated with fragile X syndrome disrupt the functioning of a gene that codes for a certain protein, designated fragile X mental retardation protein (FMRP). In normal cells, FMRP binds to dozens of different mRNA molecules (exactly why it binds is not yet understood), but in cells of individuals with fragile X syndrome the mutated allele does not produce functional FMRP.

Genomic imprinting is determined by whether inheritance is from the male or female parent Some traits are the result of genomic or parental imprinting, in which the expression of a gene in a given tissue or developmental stage is based on its parental origin—that is, whether the indi-

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deleted. As in most deletions, the exact point of breakage in chromosome 5 varies from one individual to another; some cases of cri du chat involve a small loss, whereas others involve a more substantial deletion of base pairs. Infants born with cri du chat syndrome typically have a small head with altered features described as a “moon face” and a distinctive cry that sounds like a kitten mewing. (The name literally means “cry of the cat” in French.) Affected individuals usually survive beyond childhood but exhibit severe mental retardation.

Fragile site

CGG repeats (200 to more than 1000 times)

(a) The defective allele at the tips of both chromatids of the X chromosome.

CGG repeats (up to 50 times)

(b) The normal allele.

FIGURE 16-6 Fragile X syndrome This colorized SEM shows an X chromosome with a fragile site and a normal X chromosome.

vidual inherits the gene from the male or female parent. For some imprinted genes, the paternally inherited allele is always repressed (not expressed); for other imprinted genes, the maternally inherited allele is always repressed. Thus, maternal and paternal genomes have different imprints that result in differential gene expression in the embryo. As discussed in Chapter 14, epigenetic inheritance refers to changes in how a gene is expressed without any change in the coding of the DNA bases. Epigenetic inheritance may cause genomic imprinting. Two rare genetic disorders provide a fascinating demonstration of genomic imprinting. In Prader-Willi syndrome (PWS), individuals become compulsive overeaters and obese; they are also short in stature and mildly to moderately retarded. In Angelman syndrome (AS), affected individuals are hyperactive, mentally retarded, and unable to speak, and they suffer from seizures. A small deletion of several loci from the same region of chromosome 15 causes both PWS and AS (FIG. 16-7a). One of these deleted loci is responsible for PWS, another for AS. Pedigree analysis has shown that when the person inherits the deletion from the father, PWS occurs, whereas when the person inherits the deletion from the mother, AS occurs. This inheritance pattern suggests that the normal PWS gene is expressed only in the paternal chromosome and the normal AS gene is expressed only in the maternal chromosome. PWS occurs because the repressed

KEY POINT

Many examples of genomic imprinting have been found in mammals, and more will undoubtedly be discovered.

PWS locus

Deletion includes both PWS and AS loci

AS locus

(a) Homologous chromosome 15, showing (left) close proximity of the PWS and AS genes and (right) site of deletion that includes both PWS and AS loci.

Chromosome 15 in ovum

Chromosome 15 in sperm

Repressed (imprinted) PWS gene

Chromosome 15 in ovum

Chromosome 15 in sperm

Absent AS gene

Absent PWS gene

Repressed (imprinted) AS gene

Active AS gene

From ovum

Active PWS gene

From sperm

Prader-Willi syndrome (PWS)

(b) This individual has Prader-Willi syndrome because the sperm contributed a chromosome 15 in which the PWS gene had been deleted. The chromosome with the repressed (imprinted) PWS gene from the egg is unable to compensate for the deletion.

From ovum

From sperm

Angelman syndrome (AS)

(c) This individual has Angelman syndrome because the egg contributed a chromosome 15 in which the AS gene had been deleted. The chromosome with the repressed (imprinted) AS gene from the sperm is unable to compensate for the deletion.

FIGURE 16-7 Genomic imprinting and development of Prader-Willi syndrome (PWS) and Angelman syndrome (AS) The PWS gene is active in the male gamete and repressed (imprinted) in the female gamete. Conversely, the AS gene is active in the female gamete and repressed (imprinted) in the male gamete.

(imprinted) PWS gene from the mother cannot make up for the absent PWS gene in the paternal chromosome (FIG. 16-7b). Similarly, AS occurs because the repressed (imprinted) AS gene from the father cannot make up for the absent AS gene in the maternal chromosome (FIG. 16-7c).

Review ■

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What are the specific chromosome abnormalities in Down syndrome, Klinefelter syndrome, and Turner syndrome? What is the chromosome abnormality in cri du chat syndrome? What is the chromosome abnormality in fragile X syndrome?

16.3 GENETIC DISEASES CAUSED BY SINGLE-GENE MUTATIONS ■ ■ LEARNING OBJECTIVE 6 State whether each of the following genetic defects is inherited as an autosomal recessive, autosomal dominant, or X-linked recessive: phenylketonuria (PKU), sickle cell anemia, cystic fibrosis, Tay-Sachs disease, Huntington’s disease, and hemophilia A.

You have seen that several human disorders involve chromosome abnormalities. Hundreds of human disorders, however, involve enzyme defects caused by mutations of single genes. Phenylketonuria (PKU) and alkaptonuria (discussed in Chapter 13) are examples of these disorders, each of which is sometimes referred to as an inborn error of metabolism, a metabolic disorder caused by the mutation of a gene that codes for an enzyme needed in a biochemical pathway. Both PKU and alkaptonuria involve blocks in the metabolism of specific amino acids.

Many genetic diseases are inherited as autosomal recessive traits Many human genetic diseases have a simple autosomal recessive inheritance pattern and therefore are expressed only in the homozygous state. Why are these traits recessive? Most recessive mutations result in a mutant allele that encodes a product that no longer works (either there is not enough gene product, or it is a defective gene product). In the heterozygous state, there is one functional copy of the gene and one mutated, nonfunctional copy. The normal copy of the gene generally produces enough protein to meet the cell’s needs. In homozygous recessive individuals, both alleles of the gene are nonfunctional, and the cell’s needs are not met. As a result, the person shows symptoms of disease. Phenylketonuria results from an enzyme deficiency Phenylketonuria (PKU), which is most common in individuals of western European descent, is an autosomal recessive disease caused by a defect of amino acid metabolism. It affects about 1 in 10,000 live births in North America. Homozygous recessive individuals lack an enzyme that converts the amino acid phe-

nylalanine to another amino acid, tyrosine. These individuals accumulate high levels of phenylalanine, phenylpyruvic acid, and similar compounds. The accumulating phenylalanine is converted to phenylketones, which damage the central nervous system, including the brain, in children. The ultimate result in untreated cases is severe mental retardation. An infant with PKU is usually healthy at birth because its mother, who is heterozygous, breaks down excess phenylalanine for both herself and her fetus. However, during infancy and early childhood the accumulation of toxic products eventually causes irreversible damage to the central nervous system. Beginning in the 1950s, infants with PKU have been identified early and placed on a low-phenylalanine diet, dramatically alleviating their symptoms. The diet is difficult to adhere to because it contains no meat, fish, dairy products, breads, or nuts. Also, individuals with PKU should not consume the sugar substitute aspartame, found in many diet drinks and foods, because it contains phenylalanine. Biochemical tests for PKU have been developed, and screening of newborns through a simple blood test is required in the United States. Because of these screening programs and the availability of effective treatment, thousands of PKU-diagnosed children have not developed severe mental retardation. Most must continue the diet through at least adolescence. Doctors now recommend that patients stay on the diet throughout life because some adults who have discontinued the low-phenylalanine diet experience certain mental problems, such as difficulty in concentration and short-term memory loss. Ironically, the success of PKU treatment in childhood presents a new challenge today. If a homozygous female who has discontinued the special diet becomes pregnant, the high phenylalanine levels in her blood can damage the brain of the fetus she is carrying, even though that fetus is heterozygous. Therefore, she must resume the diet, preferably before becoming pregnant. This procedure is usually (although not always) successful in preventing the effects of maternal PKU. It is especially important for women with PKU to be aware of maternal PKU and to obtain appropriate counseling and medical treatment during pregnancy. Sickle cell anemia results from a hemoglobin defect Sickle cell anemia is inherited as an autosomal recessive trait. The disease is most common in people of African descent (approximately 1 in 500 African Americans), and about 1 in 12 African Americans is heterozygous. Under low oxygen conditions, the red blood cells of an individual with sickle cell anemia are shaped like sickles, or half-moons, whereas normal red blood cells are biconcave discs. The mutation that causes sickle cell anemia was first identified in 1957. The sickled cells contain abnormal hemoglobin molecules, which have the amino acid valine instead of glutamic acid at position 6 (the sixth amino acid from the amino terminal end) in the b-globin chain (see Fig. 3-22a). The substitution of valine for glutamic acid makes the hemoglobin molecules stick to one another to form fiberlike structures that change the shape of some of the red blood cells. This sickling occurs in the veins after the oxygen has been released from the hemoglobin. The blood cells’ abnormal sickled shape slows blood flow and blocks small blood

vessels (FIG. 16-8), resulting in tissue damage from lack of oxygen and essential nutrients and in episodes of pain. Because sickled red blood cells also have a shorter lifespan than normal red blood cells, many affected individuals have severe anemia. Treatments for sickle cell anemia include pain-relief measures, transfusions, and, more recently, medicines such as hydroxyurea, which activates the gene for the production of normal fetal hemoglobin (this gene is generally not expressed after birth). The presence of normal fetal hemoglobin in the red blood cells dilutes the sickle cell hemoglobin, thereby minimizing the painful episodes and reducing the need for blood transfusions. The long-term effects of hydroxyurea are not known at this time, but there are concerns that it may induce tumor formation. Ongoing research is directed toward providing gene therapy for sickle cell anemia. The development of a mouse model for studying sickle cell anemia has enabled researchers to test gene therapy. The first gene therapy treatments in mice used a mouse retrovirus as a vector, a carrier that transfers the genetic information. However, the retrovirus did not effectively transport the normal gene for hemoglobin into the bone marrow, where stem cells produce new blood cells. In 2001, researchers cured sickle cell anemia in mice using a modified HIV as a vector. Before this treatment can be tested in humans, however, researchers must demonstrate that the HIV vector is safe. Bone marrow transplants are also a promising future treatment for seriously ill individuals. The reason the sickle cell allele occurs at a higher frequency in parts of Africa and Asia is well established. Individuals who are heterozygous (HbAHbS) and carry alleles for both normal hemoglobin (HbA) and sickle cell hemoglobin (HbS) are more resistant

to the malarial parasite, Plasmodium falciparum, which causes a severe, often fatal, form of malaria. The malarial parasite, which spends part of its life cycle inside red blood cells, does not thrive when sickle cell hemoglobin is present. (An individual heterozygous for sickle cell anemia produces both normal and sickle cell hemoglobin.) Areas in Africa where falciparum malaria occurs correlate well with areas in which the frequency of the sickle cell allele is more common in the human population. Thus, HbAHbS individuals, possessing one copy of the mutant sickle cell allele, have a selective advantage over both types of homozygous individuals, HbAHbA (who may die of malaria) and HbSHbS (who may die of sickle cell anemia). This phenomenon, known as heterozygote advantage, is discussed further in Chapter 19 (see Fig. 19-7).

Dr. Stanley Flegler/ Visuals Unlimited

Cystic fibrosis results from defective ion transport Cystic fibrosis is the most common autosomal recessive disorder in children of European descent (1 in 2500 births). About 1 in 25 individuals in the United States is a heterozygous carrier of the mutant cystic fibrosis allele. Abnormal secretions characterize this disorder. The most severe effect is on the respiratory system, where abnormally viscous mucus clogs the airways. The cilia that line the bronchi cannot easily remove the mucus, and it becomes a growth medium for dangerous bacteria. These bacteria or their toxins attack the surrounding tissues, leading to recurring pneumonia and other complications. The heavy mucus also occurs elsewhere in the body, causing digestive difficulties and other effects. As discussed earlier, the gene responsible for cystic fibrosis codes for CFTR, the protein that regulates the transport of chloride ions across cell membranes. The defective protein, found in plasma membranes of epithelial cells lining the passageways of the lungs, intestines, pancreas, liver, sweat glands, and reproductive organs, results in the production of an unusually thick mucus that eventually leads to tissue damage. Capillary Although many forms of cystic fibrosis exist that (small blood vessel) vary somewhat in the severity of symptoms, the disease is almost always serious. Sickled Antibiotics are used to control bacterial inred blood cells fections, and daily physical therapy is required to clear mucus from the respiratory system ( FIG. 16-9). Treatment with Dornase Alpha (DNase), an enzyme produced by recombinant DNA technology, helps break down the mucus. Without treatment, death would occur in infancy. With treatment, the average life expecNormal tancy for individuals with cystic fibrosis is about red blood cells 38 years. Because of the serious limitations of available treatments, gene therapy for cystic fibrosis is under development. The most severe mutant allele for cystic 10 μm fibrosis predominates in northern Europe, and another, somewhat less serious, mutant allele is more prevalent in southern Europe. Presumably, these mutant alleles are independent mutations FIGURE 16-8 Sickle cell anemia that have been maintained by natural selecSickled red blood cells do not pass through small blood vessels as easily as unsickled red blood cells do. The sickled cells can cause blockages that prevent oxygen from being delivered to tissues. tion. Some experimental evidence supports the

Hattie Young/Photo Researchers, Inc.

Some genetic diseases are inherited as autosomal dominant traits

FIGURE 16-9 Treating cystic fibrosis One traditional treatment for cystic fibrosis is chest percussion, or gentle pounding on the chest, to clear mucus from clogged airways in the lungs. This technique is typically done after nebulizer therapy, which loosens the mucus.

hypothesis that heterozygous individuals are less likely to die from infectious diseases that cause severe diarrhea, such as cholera, another possible example of heterozygote advantage.

Huntington’s disease (HD), named after George Huntington, the U.S. physician who first described it in 1872, is caused by a rare autosomal dominant allele that affects the central nervous system. The disease causes severe mental and physical deterioration, uncontrollable muscle spasms, and personality changes; death ultimately results. No effective treatment has been found. Every child of an affected individual has a 50% chance of also being affected (and, if affected, of passing the abnormal allele to his or her offspring). Ordinarily, we would expect a dominant allele with such devastating effects to occur only as a new mutation and not to be transmitted to future generations. Because HD symptoms typically do not appear until relatively late in life (most people do not develop the disease until they are in their forties), affected individuals are likely to produce children before the disease develops (FIG. 16-10). In North America, HD occurs in 1 in 20,000 live births. The gene responsible for HD is located on the short end of chromosome 4. The mutation is a nucleotide triplet (CAG) that is repeated many times; the normal allele repeats CAG from 6 to 35 times, whereas the mutant allele repeats CAG from 40 to more than 150 times. Because CAG codes for the amino acid glutamine, the resulting protein, called “huntingtin,” has a long strand of glutamines. The number of nucleotide triplet repeats seems to be important in determining the age of onset and the severity of the disease; larger numbers of repeats correlate with an earlier age of onset and greater severity. Much research now focuses on how the mutation is linked to neurodegeneration in the brain. A mouse model of HD is providing valuable clues about the development of the disease. Using this model, researchers have demonstrated that the defective version of huntingtin binds to enzymes called acetyltransferases in brain cells,

Tay-Sachs disease results from abnormal lipid metabolism in the brain Probability of developing symptoms by a given age

Tay-Sachs disease is an autosomal recessive disease that affects the central nervous system and results in blindness and severe mental retardation. The symptoms begin within the first year of life and result in death before the age of 5 years. Because of the absence of an enzyme, a normal membrane lipid in brain cells fails to break down properly and accumulates in intracellular organelles called lysosomes (discussed in Chapter 4). The lysosomes swell and cause nerve cells to malfunction. Although research is ongoing, no effective treatment for Tay-Sachs disease is available at this time. However, an effective treatment strategy in a mouse model was reported in 1997: oral administration of an inhibitor reduced the synthesis of the lipid that accumulates in the lysosomes. This treatment offers the hope of future breakthroughs to deal more effectively with Tay-Sachs disease in humans. The abnormal allele is especially common in the United States among Jewish individuals whose ancestors came from eastern and central Europe (Ashkenazi Jews). In contrast, Jewish individuals whose ancestors came from the Mediterranean region (Sephardic Jews) have a very low frequency of the allele.

1.0 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0

0

10

20

30

40

50

60

70

Age (years)

FIGURE 16-10 Age of onset of Huntington’s disease The graph shows the cumulative probability that an individual carrying a Huntington’s disease allele will have developed symptoms at a given age. (Adapted from Harper, P. S., Genetic Counseling, 5th ed., ButterworthHeinemann, Oxford, 1998.)

blocking their action. Acetyltransferases are involved in turning genes on for expression, so in the brain cells of HD individuals, much of normal transcription cannot occur. Once neurologists better understand HD’s mechanism of action on nerve cells, it may be possible to develop effective treatments to slow the progression of the disease. Cloning of the HD allele became the basis for tests that allow those at risk to learn presymptomatically if they carry the allele. The decision to be tested for any genetic disease is understandably a highly personal one. The information is, of course, invaluable for those who must decide whether or not to have children. However, someone who tests positive for the HD allele must then live with the virtual certainty of eventually developing this devastating and incurable disease. Researchers hope that information from affected individuals who choose to be identified before the onset of symptoms may ultimately contribute to the development of effective treatments.

Some genetic diseases are inherited as X-linked recessive traits Hemophilia A was once referred to as a disease of royalty because of its high incidence among male descendants of Queen Victoria, but it is also found in many nonroyal pedigrees. Caused by the absence of a blood-clotting protein called factor VIII, hemophilia A is characterized by severe internal bleeding in the head, joints, and other areas from even a slight wound. The mode of inheritance is X-linked recessive. Thus, affected individuals are almost exclusively male, having inherited the abnormal allele on the X chromosome from their heterozygous carrier mothers. (For a female to be affected by an X-linked trait, she would have to inherit the defective allele from both parents, whereas an affected male need only inherit one defective allele from his mother.) Treatments for hemophilia A consist of blood transfusions and the administration of clotting factor VIII (the missing gene product) by injection. Unfortunately, these treatments are costly. During the 1980s, many clotting factor VIII preparations made from human plasma were unknowingly contaminated with HIV, and many men with hemophilia subsequently died from AIDS. Since 1992, virus-free clotting factor VIII has been available from both human plasma and recombinant DNA technology. Geneticists are beginning to unravel X-linked genes affecting intelligence The X chromosome contains a disproportionate number of the more than 200 genes identified so far that affect cognitive abilities, for example, by coding for proteins required for the brain to function normally. Not surprisingly, many kinds of mental impairment are linked to defects in genes on the X chromosome. By one count, the human X chromosome contains less than 4% of the human genome, yet 10% of the genes in which defects are known to cause some form of mental retardation are found on the X chromosome. Because males have only one X chromosome, more boys than girls have some form of mental impairment, a fact that has been observed for more than a century.

Review ■

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Which of the following genetic diseases is/are inherited as an autosomal recessive: phenylketonuria, Huntington’s disease, Tay-Sachs disease? Which of the following genetic diseases is/are inherited as an autosomal dominant: sickle cell anemia, hemophilia A, Huntington’s disease? Which of the following genetic diseases is/are inherited as an X-linked recessive: hemophilia A, cystic fibrosis, Tay-Sachs disease?

16.4 GENE THERAPY ■ ■ LEARNING OBJECTIVE 7

Briefly discuss the process of gene therapy, including some of its technical challenges.

Because serious genetic diseases are difficult to treat, scientists have dreamed of developing actual cures. Today, advances in genetics are bringing these dreams closer to reality. One strategy is gene therapy, which aims to compensate for a defective, mutant allele by adding a normal, therapeutic allele (and its expressed protein) to certain cells. The rationale is that although a particular allele may be present in all cells, it is expressed only in some. Expression of the normal allele in only the cells that require it may be sufficient to yield a normal phenotype (FIG. 16-11). This approach presents several technical problems. The solutions to these problems must be tailored to the nature of the gene itself, as well as to its product and the types of cells in which it is expressed. First, the gene is cloned and the DNA is introduced into the appropriate cells. One of the most successful techniques is packaging the normal allele in a viral vector, a virus that moves the normal allele into target cells that currently have a mutant allele. Ideally, the virus should infect a high percentage of the cells. Most important, the virus should do no harm, especially over the long term. Early gene therapy trials used an adenovirus, which causes the common cold. However, some individuals had a strong immune reaction to the virus, so many current trials use an adeno-associated virus that does not cause side effects. To date, gene therapy has had a greater than 90% success rate in restoring the immune systems of 30 children with severe combined immunodeficiency (SCID). (SCID is a group of inherited disorders that seriously compromise the immune system.) This success rate is significantly better than the 50% success rate of the older therapy involving bone marrow transplants. Although many obstacles must be overcome, gene therapies for several other genetic diseases are under development or are being tested on individuals in clinical trials. Scientists are currently addressing some of the unique problems presented by each disease.

Gene therapy programs are carefully scrutinized Until recently, major technical advances caused the number of clinical studies involving gene therapy to grow dramatically. However, the death of a young man in a gene therapy trial in 1999, and five more recent cases of cancer (leukemia) in children, one of whom

RESEARCH METHOD Why Is It Used?

Mice are a model system for the development of gene therapy, the use of normal genes to correct or alleviate the symptoms of a genetic disease caused by defective copies of a particular gene. This procedure is performed on humans for certain types of genetic disease.

How Is It Done? Bone marrow cells extracted from mice

Virus with therapeutic gene copy

Cells placed in nutrient medium

1

Transformed cells

2

Bone marrow cells

Transformed cells

3

Cloned genes inserted into cells

4

Cloned gene 5

Transformed cells injected into recipient animal

mRNA Protein Desired protein is produced

Cells in suspension

FIGURE 16-11 Gene therapy in bone marrow cells of a mouse

died, led to a shutdown of many trials, pending the outcome of investigations about health risks. The main safety concern in these inquiries is the potential toxicity of viral vectors. The vector used in the young man who died was an adenovirus that was required in large doses to transfer enough copies of the normal alleles for effective therapy. Unfortunately, the high viral doses triggered a fatal immune response in the patient’s body. The children who developed leukemia were being treated for SCID. The vector in these cases was a retrovirus that inserted itself into and activated an oncogene that can cause childhood leukemia. Performing clinical trials on humans always has inherent risks. Researchers carefully select patients and thoroughly explain the potential benefits and risks, as far as they are known, so the patient— or, in the case of children, the parents—can give informed consent for the procedure. However, the problems in gene therapy trials in recent years have researchers busy developing safer alternatives to viral vectors.

Review ■ ■

How are viruses used in human gene therapy? Why are viral vectors of potential concern in human gene therapy?

16.5 GENETIC TESTING AND COUNSELING ■ ■ LEARNING OBJECTIVES 8

State the relative advantages and disadvantages of amniocentesis, chorionic villus sampling, and preimplantation genetic diagnosis in the prenatal diagnosis of human genetic abnormalities.

9 Distinguish between genetic screening programs for newborns and adults, and discuss the scope and implications of genetic counseling.

Geneticists have made many advances in detecting genetic disorders in individuals in recent years, including in prenatal diagnosis and genetic screening. With these advances comes increased information for couples at risk of having children with genetic diseases. Helping couples understand and deal with the genetic information now available is part of the rapidly expanding field of genetic counseling.

Prenatal diagnosis detects chromosome abnormalities and gene defects Health-care professionals are increasingly successful at diagnosing genetic diseases prenatally. In the diagnostic technique called amniocentesis, a physician obtains a sample of the amniotic fluid surrounding the fetus by inserting a needle through the pregnant woman’s abdomen, into the uterus, and then into the amniotic sac surrounding the fetus. Some of the amniotic fluid is withdrawn from the amniotic cavity into a syringe (FIG. 16-12). The fetus is normally safe from needle injuries because ultrasound imaging helps determine the positions of the fetus, placenta, and the needle. (Figure 51-18 shows an ultrasound of a human fetus.) However, there is a 0.5%, or 1 in 200, chance that amniocentesis will induce a miscarriage. Amniotic fluid contains living cells sloughed off the body of the fetus and hence genetically identical to the cells of the fetus. After cells grow for about 2 weeks in culture in the lab, technicians karyotype dividing cells to detect chromosome abnormalities. Other DNA tests have also been developed to identify most chromosome abnormalities. Amniocentesis, which has been per-

of a normally occurring protein, a-fetoprotein, in the amniotic fluid. Some of this protein crosses the placenta into the mother’s blood, which is tested for maternal serum a-fetoprotein (MSAFP) as a screen 1 About 20 mL of amniotic for spinal cord defects. If an elefluid containing cells vated level of MSAFP is detected, Ultrasound probe sloughed off from 16-week determines position of the physician performs diagnostic fetus is removed fetus fetus through mother‘s tests, such as ultrasound imaging Uterine wall abdomen. and amniocentesis. (Interestingly, Amniotic cavity abnormally low levels of MSAFP are associated with Down syndrome and other trisomies.) 2 Fluid is One problem with amniocencentrifuged. tesis is that most of the conditions it detects are unpreventable and incurable, and the results are gen3 Amniotic erally not obtained until well into fluid is the second trimester, when termianalyzed. nating the pregnancy is both psychologically and medically more 4 Fetal cells are difficult than earlier. Therefore, Placenta checked to researchers have developed tests determine sex, and that yield results earlier in the pregpurified DNA is 5 Some nancy. Chorionic villus sampling analyzed. cells are 6 Karyotype is (CVS) involves removing and grown for analyzed for sex studying cells that will form the 2 weeks in chromosomes culture medium. fetal contribution to the placenta or any chromosome (FIG. 16-13). CVS, which has been abnormality. 7 Cells are analyzed performed in the United States biochemically for since about 1983, is associated presence of about with a slightly greater risk of infec40 metabolic disorders. tion or miscarriage than amniocentesis, but its advantage is that FIGURE 16-12 Animated Amniocentesis results are obtained earlier in the Certain genetic diseases and other abnormal conditions are diagnosed prenatally by amniocentesis. pregnancy than in amniocentesis, usually within the first trimester. A relatively new embryoscreening process, known as preimplantation genetic diagnosis (PGD), is available for couples formed since the 1960s, is routinely offered for pregnant women who carry alleles for Tay-Sachs disease, hemophilia, sickle cell older than age 35 because their fetuses have a higher-than-normal anemia, and dozens of other inherited genetic conditions. PGD risk of Down syndrome. is an adjunct to assisted reproductive technology. Conception is Researchers have developed prenatal tests to detect many geby in vitro fertilization (IVF), in which gametes are collected, netic disorders with a simple inheritance pattern, but these disorders eggs are fertilized in a dish in the laboratory, and the resulting emare rare enough that physicians usually order the tests performed bryo is then implanted in the uterus for development (see Inquironly if they suspect a particular problem. Enzyme deficiencies can ing About: Novel Origins, in Chapter 50). Prior to implantation, often be detected by incubating cells recovered from amniotic fluid the physician screens single cells of early embryos for one or more with the appropriate substrate and measuring the product; this genetic diseases before placing a healthy embryo into the woman’s technique has been useful in prenatal diagnosis of disorders such uterus. PGD differs from amniocentesis and CVS in that the test as Tay-Sachs disease. The tests for several other diseases, including is performed before a woman is pregnant, so it eliminates the decisickle cell anemia, Huntington’s disease, and cystic fibrosis, involve sion of whether or not to terminate the pregnancy if an embryo directly testing the individual’s DNA for the mutant allele. has a genetic abnormality. However, PGD is not as accurate as Amniocentesis is also useful in detecting a condition known as amniocentesis or CVS, and it is more expensive. Moreover, PGD spina bifida, in which the spinal cord does not close properly during is sometimes controversial because some couples use it to choose development. A relatively common malformation (about 1 in 300 the gender of their offspring, not to screen for genetic diseases. births), this birth defect is associated with abnormally high levels

KEY POINT

In amniocentesis, the fluid surrounding the developing fetus is sampled, usually during the 16th week of pregnancy, to detect genetic and developmental disorders.

KEY POINT

In chorionic villus sampling, chorionic cells genetically identical to the embryo are sampled, usually during the eighth or ninth week of pregnancy, to detect genetic disorders.

Transabdominal sampling technique Withdrawn chorionic villi cells

Cervical sampling technique

Ultrasound probe Catheter

Syringe Withdrawn chorionic villi cells or

Chorionic Catheter villi

Cells are cultured; biochemical tests and karyotyping are performed

FIGURE 16-13 Chorionic villus sampling (CVS) This test allows the early diagnosis of some genetic abnormalities. Samples may be obtained by inserting a needle through the uterine wall or the cervical opening.

Although using amniocentesis, CVS, and PGD can help physicians diagnose certain genetic disorders with a high degree of accuracy, the tests are not foolproof, and many disorders cannot be diagnosed at all. Therefore, the lack of an abnormal finding is no guarantee of a normal baby.

Genetic screening searches for genotypes or karyotypes Genetic screening is a systematic search through a population for individuals with a genotype or karyotype that might cause a serious genetic disease in themselves or their offspring. There are two main types of genetic screening, for newborns and for adults, and each serves a different purpose. Newborns are screened primarily as the first step in preventive medicine, and adults are screened to help them make informed reproductive decisions. Newborns are screened to detect and treat certain genetic diseases before the onset of serious symptoms. The routine screening of infants for PKU began in 1962 in Massachusetts. Laws in all 50 states of the United States, as well as in many other countries, currently require PKU screening. Sickle cell anemia is also more effectively treated with early diagnosis. Screening newborns for sickle

cell anemia reduces infant mortality by about 15% because doctors can administer daily doses of antibiotics, thereby preventing bacterial infections common to newborns with the disease. The number of genetic disorders that can be screened in newborns is rapidly increasing, and the March of Dimes currently recommends screening newborns for 29 disorders, most of which are genetic. Genetic screening of adults identifies carriers (heterozygotes) of recessive genetic disorders. If both prospective parents are heterozygous, the carriers are counseled about the risks involved in having children. Since the 1970s, about 1 million young Jewish adults in the United States, Israel, and other countries have been screened voluntarily for Tay-Sachs disease, and about 1 in 30 has been identified as a carrier. Tay-Sachs screening programs have reduced the incidence of TaySachs disease to almost zero.

Genetic counselors educate people about genetic diseases

Couples who are concerned about the risk of abnormality in their children, because they have either an abnormal child or a relative affected by a hereditary disease, may seek genetic counseling for medical and genetic information as well as support and guidance. Genetic clinics, available in most major metropolitan centers, are usually affiliated with medical schools. Genetic counselors, who have received training in counseling, medicine, and human genetics, provide people with the information they need to make reproductive decisions. They offer advice, tempered with respect and sensitivity, in terms of risk estimates— that is, the probability that any given offspring will inherit a particular condition. The counselor uses the complete family histories of both the man and the woman, and a clinical geneticist (a physician who specializes in genetics) may screen for the detection of heterozygous carriers of certain conditions. When a disease involves only a single gene locus, probabilities can usually be easily calculated. For example, if one prospective parent is affected with a trait that is inherited as an autosomal dominant disorder, such as Huntington’s disease, the probability that any given child will have the disease is 0.5, or 50%. The birth to phenotypically normal parents of a child affected with an autosomal recessive trait, such as albinism or PKU, establishes that both parents are heterozygous carriers, and the probability that any subsequent child will be affected is therefore 0.25, or 25%. For a disease inherited through a recessive allele on the X chromo-

some, such as hemophilia A, a normal woman and an affected man will have daughters who are carriers and sons who are normal. The probability that the son of a carrier mother and a normal father will be affected is 0.5, or 50%; the probability that their daughter will be a carrier is also 0.5, or 50%. It is important for identified carriers to receive appropriate genetic counseling. A genetic counselor is trained not only to provide information pertaining to reproductive decisions but also to help individuals understand their situation and avoid feeling stigmatized.

Review ■

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What are the relative advantages and disadvantages of amniocentesis, chorionic villus sampling, and preimplantation genetic diagnosis? What is the purpose of genetic screening for newborns? What is the purpose of genetic screening for adults?

16.6 HUMAN GENETICS, SOCIETY, AND ETHICS ■ ■ LEARNING OBJECTIVE 10 Discuss the controversies of genetic discrimination.

Many misconceptions exist about genetic diseases and their effects on society. Some people erroneously think of certain individuals or populations as genetically unfit and thus responsible for many of society’s ills. They argue, for example, that medical treatment of people affected with genetic diseases, especially those who reproduce, increases the frequency of abnormal alleles in the population. However, such notions are incorrect. Genetic disorders are so rare that modern medical treatments will have only a negligible effect on their incidence. Recessive mutant alleles are present in all individuals and all ethnic groups; no one is exempt. According to one estimate, each of us is heterozygous for several (3 to 15) harmful recessive alleles, any of which could cause debilitating illness or death in the homozygous state. Why, then, are genetic diseases relatively uncommon? Each of us has many thousands of essential genes, any of which can be mutated. It is very unlikely that the abnormal alleles that one individual carries are also carried by that individual’s mate. Of course, this possibility is more likely if the harmful allele is a relatively common one, such as the one responsible for cystic fibrosis. Relatives are more likely than nonrelatives to carry the same harmful alleles, having inherited them from a common ancestor. A greater-than-normal frequency of a particular genetic disease among offspring of consanguineous matings, matings between genetically related individuals, is often the first clue that the mode of inheritance is autosomal recessive. The offspring of consanguineous matings have a small but significantly increased risk of genetic disease. In fact, they account for a disproportionately high percentage of those individuals in the population with autosomal recessive disorders. Because of this perceived social cost, marriages of close relatives, including first cousins, are prohibited by about half the

states in the United States. However, consanguineous marriages are still relatively common in many other countries.

Genetic discrimination provokes heated debate One of the fastest-growing areas of medical diagnostics is genetic screening and testing, and new genetic tests that screen for diseases such as cystic fibrosis, sickle cell anemia, Huntington’s disease, colon cancer, and breast cancer increase each year. However, genetic testing raises many social, ethical, and legal issues that we as a society must address. One of the most difficult issues is whether genetic information should be available to health insurance and life insurance companies. Many people think genetic information should not be given to insurance companies, but others, including employers, insurers, and many organizations representing people affected by genetic disorders, say such a view is unrealistic. If people use genetic tests to help them decide when to buy insurance and how much, then insurers insist they should also have access to this information. Insurers say they need access to genetic data to help calculate equitable premiums (insurance companies average risks over a large population). However, some are concerned that insurers might use the results of genetic tests to discriminate against people with genetic diseases or to deny them coverage. Physicians argue that people at risk for a particular genetic disease might delay being tested because they fear genetic discrimination from insurers and employers. Genetic discrimination is discrimination against an individual or family members because of differences from the “normal” genome in that individual. The perception of genetic discrimination already exists in society. A 1996 study found that 25% of 332 people with family histories of one or more genetic disorders thought they had been refused life insurance, 22% thought they had been refused health insurance, and 13% thought they had been denied employment because of genetic discrimination. In a 1998 survey by the National Center for Genetic Resources, 63% of respondents said they probably or definitely would not take a genetic test if the results could be disclosed to either their employers or insurers. Complicating the issue even more, genetic tests are sometimes difficult to interpret, in part because of the many complex interactions between genes and the environment. If a woman tests positive for an allele that has been linked to breast cancer, for example, she is at significant risk, but testing positive does not necessarily mean she will develop breast cancer. These uncertainties also make it hard to decide what form of medical intervention, from frequent mammograms to surgical removal of healthy breasts, is appropriate. The Ethical, Legal, and Social Implications (ELSI) Research Program of the National Human Genome Research Institute has developed principles designed to protect individuals against genetic discrimination. In 2008 the Genetic Information Nondiscrimination Act (GINA) extended significant protection against workplace and health discrimination. This law prohibits employers and insurance companies from discriminating on the basis of information derived from genetic tests. For example, neither employers nor insurance companies are permitted to request a genetic test. Employers cannot base employment decisions on

genetic information. Also, insurance companies cannot cancel or deny coverage or increase the price of premiums based on genetic information.

Many ethical issues related to human genetics must be addressed Genetic discrimination is only one example of ethical issues arising from our expanding knowledge of human genetics. Consider the following questions, all of which deal with the broad ethical issue of individual rights: What is the youngest age at which genetic testing should be permitted for adult-onset diseases, such as Huntington’s disease? What are the emotional and psychological effects on individuals who are told they have tested positive for an incurable genetic disease? Should testing be performed when some family members want testing and others do not? Should parents be able to test their minor children? Should access to genetic test data be

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permitted in cases of paternity or kinship testing? Should states be able to collect genetic data on their residents? Should school administrators or law enforcement agencies have access to genetic data? These questions are only a sample of the many issues that both ethicists and society must consider now and in the future. As human genetics assumes an increasingly important role in society, issues of genetic privacy and the confidentiality of genetic information must be addressed.

Review Why is it incorrect to assume that certain individuals or populations carry most of the abnormal alleles found in humans? To be expressed, an autosomal recessive genetic disease must be homozygous. What relationship does this fact have to consanguineous matings? Why do health and life insurance companies want genetic information about their clients?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

16.1 (page 348)

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1 Distinguish between karyotyping and pedigree analysis. ■

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Studies of an individual’s karyotype, the number and kinds of chromosomes present in the nucleus, enable researchers to identify various chromosome abnormalities. A pedigree is a “family tree” that shows the transmission of genetic traits within a family over several generations. Pedigree analysis is useful in detecting autosomal dominant mutations, autosomal recessive mutations, X-linked recessive mutations, and defects due to genomic imprinting, which is the expression of a gene based on its parental origin. Learn more about pedigrees by clicking on the figure in CengageNOW.

2 Discuss the implications of the Human Genome Project. The human genome is the total genetic information in human cells. The Human Genome Project sequenced the DNA in the nuclear human genome. Now that the human genome has been sequenced, researchers will be busy for decades identifying genes and understanding what each gene’s role is, how each gene interacts with other genes, and how the expression of each gene is regulated in different tissues. For example, researchers have begun to use genome-wide association (GWA) scans to identify genetic variations associated with complex human diseases. 3 Discuss the mouse model for studying cystic fibrosis. ■ The use of animal models greatly helps researchers investigate human disease. Researchers used gene targeting to produce strains of mice that are either homozygous or heterozygous for cystic fibrosis. Results from these studies may yield more effective drugs for treating the disease. ■

16.2 (page 351) 4 Explain how nondisjunction in meiosis is responsible for chromosome abnormalities such as Down syndrome, Klinefelter syndrome, and Turner syndrome. ■ In aneuploidy, there are either missing or extra copies of certain chromosomes. Aneuploidies include trisomy, in which an individual’s cells contain an extra chromosome, and monosomy, in which one member of a pair of chromosomes is missing.

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Trisomy 21, the most common form of Down syndrome, and Klinefelter syndrome (XXY) are examples of trisomy. Turner syndrome (X0) is an example of monosomy. Trisomy and monosomy are caused by meiotic nondisjunction, in which sister chromatids or homologous chromosomes fail to move apart properly during meiosis. Learn more about karyotypes by clicking on the figure in CengageNOW.

5 Distinguish among the following structural abnormalities in chromosomes: translocations, deletions, and fragile sites. ■ In a translocation, part of one chromosome becomes attached to another. About 4% of individuals with Down syndrome have a translocation in which the long arm of chromosome 21 is attached to the long arm of one of the larger chromosomes, such as chromosome 14. ■ A deletion can result in chromosome breaks that fail to rejoin. The deletion may range in size from a few base pairs to an entire chromosome arm. One deletion disorder in humans is cri du chat syndrome, in which part of the short arm of chromosome 5 is deleted. A

A

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Fragile sites may occur at specific locations on both chromatids of a chromosome. In fragile X syndrome, a fragile site occurs near the tip on the X chromosome, where the nucleotide triplet CGG is repeated many more times than is normal. Fragile X syndrome is the most common cause of inherited mental retardation.

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Learn more about amniocentesis by clicking on the figure in CengageNOW.

16.3 (page 358) 6 State whether each of the following genetic defects is inherited as an autosomal recessive, autosomal dominant, or X-linked recessive: phenylketonuria (PKU), sickle cell anemia, cystic fibrosis, Tay-Sachs disease, Huntington’s disease, and hemophilia A. ■ Most human genetic diseases that show a simple inheritance pattern are transmitted as autosomal recessive traits. Phenylketonuria (PKU) is an autosomal recessive disorder in which toxic phenylketones damage the developing nervous system. Sickle cell anemia is an autosomal recessive disorder in which abnormal hemoglobin (the protein that transports oxygen in the blood) is produced. Cystic fibrosis is an autosomal recessive disorder in which abnormal secretions are produced primarily in organs of the respiratory and digestive systems. Tay-Sachs disease is an autosomal recessive disorder caused by abnormal lipid metabolism in the brain. ■ Huntington’s disease is an autosomal dominant inheritance pattern that results in mental and physical deterioration, usually beginning in adulthood. ■ Hemophilia A is an X-linked recessive disorder that results in a defect in a blood component required for clotting.

16.4 (page 361) 7 Briefly discuss the process of gene therapy, including some of its technical challenges. ■ In gene therapy, the normal allele is cloned and the DNA is introduced into certain human cells where its expression may be sufficient to yield a normal phenotype. ■ One technical challenge in gene therapy is finding a safe, effective vector, usually a virus, to deliver the gene of interest into the cells.

16.5 (page 362) 8 State the relative advantages and disadvantages of amniocentesis, chorionic villus sampling, and preimplantation genetic diagnosis in the prenatal diagnosis of human genetic abnormalities.

In amniocentesis, a physician samples the amniotic fluid surrounding the fetus and cultures and screens the fetal cells suspended in the fluid for genetic defects. Amniocentesis provides results in the second trimester of pregnancy.

In chorionic villus sampling (CVS), a physician removes and studies some of the fetal cells. CVS provides results in the first trimester of pregnancy but is associated with a slightly greater risk of infection and miscarriage than amniocentesis. ■ Couples who conceive by in vitro fertilization may elect to have preimplantation genetic diagnosis (PGD). A physician screens the embryos for one or more genetic diseases before placing a healthy embryo into the woman’s uterus. PGD is not as accurate as amniocentesis or CVS, and it is more expensive. 9 Distinguish between genetic screening programs for newborns and adults, and discuss the scope and implications of genetic counseling. ■ Genetic screening identifies individuals who might carry a serious genetic disease. Screening of newborns is the first step in preventive medicine, and screening of adults helps them make informed reproductive decisions. ■ Couples who are concerned about the risk of abnormality in their children may seek genetic counseling. A genetic counselor provides medical and genetic information pertaining to reproductive decisions and helps individuals understand their situation and avoid feeling stigmatized. ■

16.6 (page 365) 10 Discuss the controversies of genetic discrimination. ■

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Genetic discrimination is discrimination against an individual or family member because of differences from the “normal” genome in that individual. One of the most difficult issues in avoiding genetic discrimination is whether genetic information should be available to employers and to health and life insurance companies. The Genetic Information Nondiscrimination Act (GINA) prohibits employers and insurance companies from discriminating on the basis of information derived from genetic tests.

T E S T YO U R U N D E R S TA N D I N G 1. A diagram of a pedigree shows (a) controlled matings between members of different true-breeding strains (b) the total genetic information in human cells (c) a comparison of DNA sequences among genomes of humans and other species (d) the subtle genetic differences among unrelated people (e) the expression of genetic traits in the members of two or more generations of a family 2. Which pattern of inheritance is associated with a trait that (1) is not usually expressed in the parents, (2) is expressed in about one-fourth of the children, and (3) is expressed in both male and female children? (a) autosomal recessive (b) autosomal dominant (c) X-linked recessive (d) X-linked dominant (e) Y-linked 3. An abnormality in which there is one more or one fewer than the normal number of chromosomes is called (a) a karyotype (b) a fragile site (c) an aneuploidy (d) trisomy (e) a translocation

4. The failure of chromosomes to separate normally during cell division is called (a) a fragile site (b) an inborn error of metabolism (c) a satellite knob (d) a translocation (e) nondisjunction 5. The chromosome composition of an individual or cell is called its (a) karyotype (b) nucleotide triplet repeat (c) pedigree (d) DNA microarray (e) translocation 6. An inherited disorder caused by a defective or absent enzyme is called (a) a karyotype (b) trisomy (c) a reciprocal translocation (d) an inborn error of metabolism (e) an aneuploidy 7. In _____, a genetic mutation codes for an abnormal hemoglobin molecule that is less soluble than usual and more likely than normal to deform the shape of the red blood cell. (a) Down syndrome (b) Tay-Sachs disease (c) sickle cell anemia (d) PKU (e) hemophilia A

8. During this procedure, a sample of the fluid that surrounds the fetus is obtained by inserting a needle through the walls of the abdomen and uterus. (a) DNA marking (b) chorionic villus sampling (c) ultrasound imaging (d) preimplantation genetic diagnosis (e) amniocentesis

9–11. Examine the following pedigrees and decide on the most likely mode of inheritance of each disorder. (a) autosomal recessive (b) autosomal dominant (c) X-linked recessive (d) a, b, or c (e) a or c 9.

10.

11.

CRITICAL THINKING 1. SCIENCE, TECHNOLOGY, AND SOCIETY. Imagine that you are a genetic counselor. What advice or suggestions might you give in the following situations? a. A couple has come for advice because the woman had a sister who died of Tay-Sachs disease. b. A young man and woman who are not related are engaged to be married. However, they have learned that the man’s parents are first cousins, and they are worried about the possibility of increased risk of genetic defects in their own children. c. A young woman’s paternal uncle (her father’s brother) has hemophilia A. Her father is free of the disease, and there has never been a case of hemophilia A in her mother’s family. Should she be concerned about the possibility of hemophilia A in her own children? d. A 20-year-old man is seeking counseling because his father was recently diagnosed with Huntington’s disease. e. A 45-year-old woman has just been diagnosed with Huntington’s disease. She says she will not tell her college-age sons because of the burden it will place on them. Given that the

woman, not her sons, is your client, do you have a duty to inform the sons? Explain your reasoning. 2. A common belief about human genetics is that an individual’s genes alone determine his or her destiny. Explain why this is a misconception. 3. Is a chromosome deletion equivalent to a frameshift mutation (discussed in Chapter 13)? Why or why not? 4. EVOLUTION LINK. Explain some of the evolutionary implications that one can conclude from the fact that mice and humans have about 500 DNA segments that are completely identical. 5. ANALYZING DATA. Examine Figure 16-10, and estimate the age at which half of individuals carrying a Huntington’s disease allele will have developed symptoms. At what age will threefourths of these individuals have symptoms? Additional questions are available in CengageNOW at www.cengage.com/ login.

17

Developmental Genetics

Stephen W. Paddock

Differential gene expression in the model organism Drosophila melanogaster.

KEY CONCEPTS

D

17.1 Differential gene expression controls the develop-

biologists knew little about how genes interact to control development.

ment of specialized tissues and organs.

Unraveling the genetic interactions that take place during development

17.2 Many genes that regulate development are quite similar in a wide range of organisms, from fruit flies to humans. Mutations in genes that regulate development have provided insights into how those genes function.

was an intractable problem using traditional methods. However, rapid

17.3 Mutations in oncogenes and tumor suppressor genes may lead to cancer.

evelopmental genetics is the study of the genes involved in cell differentiation and development of an organism. Until the late 1970s,

progress in recombinant DNA research led scientists to search for developmental mutants and to apply sophisticated techniques to study them. (A mutant is an individual with an abnormal phenotype caused by a gene mutation.) The organism in the photograph is a developing embryo of the fruit fly Drosophila melanogaster. Geneticists used immunofluorescence, in which a fluorescent dye is joined to an antibody that binds to a specific protein to localize the protein. In this case researchers bound three different antibodies—one red, one blue, and one yellow—to three specific proteins. The patterns of colored bands indicate that different cells of the embryo have differential gene expression—that is, different genes are active at the same time. Work with Drosophila and other organisms has profound implications for understanding both normal human development (including aging) and malfunctions that lead to birth defects and cancer. Striking similarities among genes that govern development in widely different species suggest that developmentally important genetic mechanisms are deeply rooted in the evolutionary history of multicellular organisms. There are

also differences in species’ developmental patterns that reflect their separate evolutionary paths. As you will learn in Chapter 18, developmental genes have played a role in reshaping organisms during the course of their evolution. Biologists are now studying how genes are activated, inactivated, and modified to control development. Eventually, scientists expect to understand how a single cell—a fertilized egg—develops into a multicellular organism as complex as a human.

17.1 CELL DIFFERENTIATION AND NUCLEAR EQUIVALENCE ■ ■ LEARNING OBJECTIVES 1 2 3

Distinguish between cell determination and cell differentiation, and between nuclear equivalence and totipotency. Describe the classic experiments of Steward, Gurdon, and Wilmut. Define stem cells, distinguish between embryonic stem cells and pluripotent stem cells, and describe some of the promising areas of research involving stem cells.

The study of development, broadly defined as all the changes that occur in the life of an individual, encompasses some of the most fascinating and difficult problems in biology today. Of particular interest is the process by which cells specialize and organize into a complex organism. During the many cell divisions required for a single cell to develop into a multicellular organism, groups of cells become gradually committed to specific patterns of gene activity through the process of cell determination. As cell determination proceeds, it restricts an embryonic cell’s developmental pathway so that its fate becomes more and more limited. The final step leading to cell specialization is cell differentiation. A differentiated cell, which has a characteristic appearance and characteristic activities, appears to be irreversibly committed to its fate. Another intriguing part of the developmental puzzle is the building of the body. In morphogenesis, the development of form, cells in specific locations differentiate and become spatially organized into recognizable structures. Morphogenesis proceeds through the multistep process of pattern formation, the organization of cells into three-dimensional structures. Pattern formation includes signaling between cells, changes in cell shapes, and cell migrations. Depending on their location, cells are exposed to different concentrations of signaling molecules that specify positional information. Thus, where a given cell is located often determines what it will become when it matures. The human body, like that of other vertebrates, contains about 250 recognizably different types of cells (FIG. 17-1). Combinations of these specialized cells, known as differentiated cells, are organized into diverse and complex structures—such as the eye, hand, and brain—each capable of carrying out many sophisticated activities. Most remarkable of all is the fact that all the structures of the body and the different cells within them descend from a unicellular zygote, a fertilized egg.

All multicellular organisms undergo complex patterns of development. The root cells of plants, for example, have structures and functions very different from those of the various types of cells located in leaves. Diversity is also found at the molecular level; most strikingly, each type of plant or animal cell makes a highly specific set of proteins. In some cases, such as the protein hemoglobin in red blood cells, one cell-specific protein may make up more than 90% of the cell’s total mass of protein. Other cells may have a complement of many cell-specific proteins, each of which is present in small amounts but still plays an essential role. However, because certain proteins are required in every type of cell (all cells, for example, require the same enzymes for glycolysis), cell-specific proteins usually make up only a fraction of the total number of different kinds of proteins. When researchers first discovered that each type of differentiated cell makes a unique set of proteins, some scientists hypothesized that each group of cells loses the genes it does not need and retains only those required. However, this does not generally seem true. According to the principle of nuclear equivalence, the nuclei of essentially all differentiated adult cells of an individual are genetically (though not necessarily metabolically) identical to one another and to the nucleus of the zygote from which they descended. This means that virtually all somatic cells in an adult have the same genes. However, different cells express different subsets of these genes. Somatic cells are all the cells of the body other than germ line cells, which ultimately give rise to a new generation. In animals, germ line cells—whose descendants ultimately undergo meiosis and differentiate into gametes—are generally set aside early in development. In plants, the difference between somatic cells and germ line cells is not as distinct, and the determination that certain cells undergo meiosis is made much later in development. The evidence for nuclear equivalence comes from cases in which differentiated cells or their nuclei have been found to retain the potential of directing the development of the entire organism. Such cells or nuclei are said to exhibit totipotency.

Most cell differences are due to differential gene expression Because genes do not seem to be lost regularly during development (and thus nuclear equivalence is present in different cell types), differences in the molecular composition of cells must be regulated by the activities of different genes. The process of developmental gene regulation is often referred to as differential gene expression. As discussed in Chapter 14, the expression of eukaryotic genes is regulated in many ways and at many levels. For example, a particular enzyme may be produced in an inactive form and then be activated later. However, much of the regulation that is important in development occurs at the transcriptional level. The transcription of certain sets of genes is repressed, whereas that of other sets is activated. Even the expression of genes that are constitutive— that is, constantly transcribed—is regulated during development so that the quantity of each product varies from one tissue type to another.

KEY POINT

As development proceeds, somatic cells that previously had the potential to develop into a variety of cells become increasingly committed to a specific fate. Red blood cell Endothelial cells

Pigment cell

Blood Heart, blood vessels

Smooth muscle cells

Adrenal gland (cortex)

Gut muscle

Retinal cell

Striated muscle cell

Tubule cell Kidney

Neuron Nervous system

Skeletal muscles

Gonad

Adrenal gland (medulla)

Cartilage cells

Mammary gland cell Glands

Bones and connective tissue

Exocrine cell Liver Pancreas

Lungs

Urinary bladder lining

Epithelial cell

Epidermis

Mesoderm Ectoderm

Skin cell Endoderm

Tracheal cell Somatic cells

Pharynx lining

Germ line Thyroid gland

Gamete-producing cells

Cell division and cell differentiation Fertilized egg

FIGURE 17-1 Vertebrate cell lineages Repeated divisions of the fertilized egg (bottom) result in the establishment of tissues containing groups of specialized cells. Germ line cells (cells that produce the gametes) are set aside early in development.

We can think of differentiation as a series of pathways leading from a single cell to cells in each of the different specialized tissues, arranged in an appropriate pattern. At times a cell makes genetic commitments to the developmental path its descendants will follow. These commitments gradually restrict the development of the descendants to a limited set of final tissue types. Determination, then, is a progressive fixation of the fate of a cell’s descendants.

Somatic cells progress along various developmental pathways undergoing a series of commitments that progressively determine their fates.

As the development of a cell becomes determined along a differentiation pathway, its physical appearance may or may not change significantly. Nevertheless, when a stage of determination is complete, the changes in the cell usually become self-perpetuating and are not easily reversed. Cell differentiation is usually the last stage in the developmental process. At this stage, a precursor cell becomes structurally and functionally recognizable as a bone cell,

for example, and its pattern of gene activity differs from that of a neuron (nerve cell) or any other cell type.

KEY EXPERIMENT QUESTION: Are differentiated somatic plant cells totipotent?

A totipotent nucleus contains all the instructions for development In plants, some differentiated cells can be induced to become the equivalent of embryonic cells. Biologists use tissue culture techniques to isolate individual cells from certain plants and to allow them to grow in a nutrient medium. In the 1950s, F. C. Steward and his coworkers at Cornell University conducted some of the first experiments investigating cell totipotency in plants (FIG. 17-2). Totipotent cells have the potential to give rise to all parts of an organism because they contain a complete set of genetic instructions required to direct the normal development of an entire organism. Steward and his colleagues induced root cells from a carrot to divide in a liquid nutrient medium, forming groups of cells called embryoid (embryo-like) bodies. These clumps of dividing cells were then transferred to an agar medium, which provided nutrients and a solid supporting structure for the developing plant cells. Some of the cells of the embryoid bodies gave rise to roots, stems, and leaves. The resulting small plants, called plantlets to distinguish them from true seedlings, were then transplanted to soil, where they ultimately developed into adult plants capable of producing flowers and viable seeds. Because these plants are all derived from the same parent plant, they are genetically alike and therefore constitute a clone. As mentioned in Chapter 15, a clone consists of individual organisms, cells, or DNA molecules that are genetically identical to another individual, cell, or DNA molecule, from which it was derived. The methods of plant tissue culture are now extensively used to produce genetically engineered plants because they enable researchers to regenerate whole plants from individual cells that have incorporated recombinant DNA molecules (see Chapter 38). In the 1950s, researchers began testing whether steps in the process of determination are reversible in animal cells by transplanting the nucleus of a cell in a relatively late stage of development into an egg cell that had been enucleated (that is, its own nucleus had been destroyed). Robert Briggs and Thomas J. King of the Institute for Cancer Research in Pennsylvania pioneered nuclear transplantation experiments. They transplanted nuclei from frog cells at different stages of development into egg cells whose nuclei had been removed. Some of the transplants proceeded normally through several developmental stages, and a few even developed into normal tadpoles. As a rule, the nuclei transplanted from cells at earlier stages were most likely to support development to the tadpole stage. As the fate of the cells became more and more determined, the probability quickly declined that a transplanted nucleus could control normal development. British biologist John B. Gurdon carried out experiments on nuclear transplantation in frogs during the 1960s. In a few cases he demonstrated that nuclei isolated from the intestinal epithelial cells of a tadpole directed development up to the tadpole stage (FIG. 17-3). This result occurred infrequently (about 1.5% of the time); however, in these kinds of experiments success counts more

HYPOTHESIS: Differentiated somatic carrot cells can be induced to develop into an entire plant. EXPERIMENT: F. C. Steward and his coworkers cultured carrot root tissues in a liquid nutrient medium. These cells divided to form clumps of undifferentiated cells. The clumps were then transferred to a solid growth medium. Mature plant

3

When transferred to soil, plantlets grow into mature plants.

2

1

Carrot root tissues are cut into discs and cultured in a liquid nutrient medium. Cells divide, forming mass of undifferentiated cells.

If transferred to solid medium, some of clumps form tiny plantlets.

RESULTS AND CONCLUSION: The development of a complete carrot plant from differentiated somatic cells demonstrated the totipotency of these cells. Source: Shantz, E. M., and F. C. Steward. “Investigations on Growth and Metabolism of Plant Cells VII: Sources of Nitrogen for Tissue Cultures under Optimal Conditions for Their Growth.” Annals of Botany, Vol. 23, 371–390, 1959. By permission of Oxford University Press.

FIGURE 17-2 Steward’s experiment on cell totipotency

in carrots

than failure. Therefore, he could safely conclude that at least some nuclei of differentiated animal cells are in fact totipotent. For many years, because these successes with frogs could not be repeated with mammalian embryos, many developmental biologists concluded that some fundamental feature of mammalian reproductive biology might be an impenetrable barrier to mammalian cloning. This perception changed markedly in 1996 and 1997 with the first reports of the birth of cloned mammals.

KEY EXPERIMENT QUESTION: Are nuclei in differentiated animal cells totipotent? HYPOTHESIS: Nuclei from differentiated cells contain the information required for normal development. EXPERIMENT: John Gurdon injected the nuclei of differentiated cells (tadpole intestinal cells) into eggs whose own nuclei were destroyed by ultraviolet radiation. Tadpole

Remove nucleus Intestinal cells Success rate: 1.5%

Inject nucleus into egg Irradiate with UV light to kill nucleus Unfertilized egg

Blastula

RESULTS AND CONCLUSION: Normal development proceeded to the tadpole stage in about 1.5% of trials, indicating that the genes for programming development up to that point were still present and could be appropriately activated. Source: Gurdon, J. B. “The Developmental Capacity of Nuclei Taken from Intestinal Epithelium Cells of Feeding Tadpoles.” Journal of Embryology and Experimental Morphology Vol. 10, Dec. 1962.

FIGURE 17-3 Gurdon’s experiment on nuclear totipotency in frogs

The first cloned mammal was a sheep In 1996, Ian Wilmut, Keith Campbell, and their coworkers at the Roslin Institute in Edinburgh, Scotland, reported that they had succeeded in cloning sheep by using nuclei from an early stage of sheep embryos (the blastocyst stage; see Chapter 51). These scientists received worldwide attention in early 1997 when they announced the birth of a lamb named Dolly (after the singer Dolly Parton). Dolly’s genetic material was derived from a cultured sheep mammary gland cell that was fused with an enucleated sheep’s egg. The resulting cell divided and developed into an embryo that was then cultured in vitro until it reached a stage at which it could be transferred to a host mother (FIG. 17-4). Not surprisingly, the overall success rate was low: of 277 fused cells, only 29 developed into embryos that could be transferred, and Dolly was the only live lamb produced. Why did Wilmut’s team succeed when so many other researchers had failed? Applying the basic principles of cell biology, they recognized that the cell cycles (see Chapter 10) of the egg cytoplasm and the donor nucleus were not synchronous. The egg cell is arrested at metaphase II of meiosis, whereas the actively growing donor somatic cell is usually in the DNA synthesis phase (S), or in G2. By withholding certain nutrients from the mammary gland cells used as donors, the researchers caused these cells to enter a nondividing state referred to as G0. This had the effect of synchronizing the cell cycles of the donor nucleus and the egg. They then used an electric shock to fuse the donor cell with the egg and initiate embryo development. Although an extremely high level of technical expertise is required, these and other researchers have modified and extended these techniques to produce cloned calves, pigs, horses, rats, mice, dogs, and cats, among others. The list of mammalian species suc-

cessfully cloned continues to grow. However, the success rate for each set of trials is low, around 1% to 2%, and the incidence of genetic defects is high. Dolly was euthanized at age 6 because she was suffering from a virus-induced lung cancer that infected several sheep where she was housed. However, she developed arthritis at 5½ years, which is relatively young for a sheep to have this degenerative disease. Some biologists speculate that using adult genetic material to produce a clone might produce an animal with prematurely old cells (see discussion of telomeres and cell aging in Chapter 12). Further research may provide some answers to this potential problem. The main focus of cloning research is the production of transgenic organisms, in which foreign genes have been incorporated (see Chapter 15). Researchers are actively pursuing new techniques to improve the efficiency of the cloning process. Only then will it be possible to produce large numbers of cloned transgenic animals for a variety of uses, such as increasing the populations of endangered species. For example, the first healthy clone of an endangered species, a wild relative of cattle known as a banteng, was born in 2003. The nucleus for this clone came from a frozen skin cell of a banteng that died in 1980 at the San Diego Zoo.

Stem cells divide and give rise to differentiated cells Stem cells are undifferentiated cells that can divide to produce differentiated descendants yet retain the ability to divide to maintain the stem cell population. When a stem cell divides, its daughter cells can remain stem cells or differentiate into specialized cells such as muscle cells, neurons, or blood cells. What happens depends on the presence or absence of an array of biochemical signals. One of

KEY EXPERIMENT QUESTION: Are nuclei in differentiated mammalian cells totipotent?

HYPOTHESIS: The nucleus of a differentiated cell from an adult mammal fused with an enucleated egg can provide the genetic information to direct normal development. EXPERIMENT: Ian Wilmut and his colleagues produced a sheep embryo by fusing a cultured adult sheep mammary cell with an enucleated sheep’s egg. He then implanted the embryo into the uterus of a host mother. Sheep mammary cell

Enucleated egg cell

Electrical shock Fusion

Egg cytoplasm containing donor cell nucleus

Embryo

Transfer to host mother

Lamb

RESULTS AND CONCLUSION: Normal development proceeded, and a female lamb—the world’s first cloned mammal—was born. To provide additional evidence that the cloned sheep was fully functioning, when she matured, she was bred and gave birth to a normal offspring. Source: Campbell, K. H., J. McWhir, W. A. Ritchie, and I. Wilmut. “Sheep Cloned by Nuclear Transfer from a Cultured Cell Line.” Nature, Vol. 380, 1996.

FIGURE 17-4 Animated Wilmut’s experiment on

mammalian cloning

the most challenging areas of stem cell research today involves determining the identity, order, and amounts of the chemical signals that will result in a specific type of cell differentiation. The most versatile stem cells—zygotes—are totipotent and have the potential to give rise to all tissues of the body and placenta. Stem cells can also be derived from embryos or adult cells. (The term adult is somewhat misleading because adult stem cells can be harvested from umbilical cord tissue, infants, and children as well as adults.) Embryonic and adult stem cells are known as pluripotent stem cells because they can give rise to many, but not all, of the types of cells in an organism. Embryonic stem cells (ES cells), formed after a zygote has undergone several rounds of cell division to form a 5- or 6-day-old blastocyst, are more versatile than adult stem cells (FIG. 17-5). For example, ES cells are pluripotent and have the potential to develop into any type of cell in the body; ES cells are not totipotent because they cannot form cells of the placenta. Adult stem cells have been found in the human brain, retina, heart, bone marrow, dental pulp, intestines, and other sites. Adult neural stem cells in the brain are pluripotent and differentiate to form neurons and glial cells. Adult stem cells in the bone marrow form both red blood cells and the various types of white blood cells of the immune system. The range of cells that adult stem cells can differentiate into is more limited than that of ES cells. However, recent studies suggest that even specialized stem cells may be more versatile than once thought. For example, adult neural stem cells form blood cells when transplanted into bone marrow. Stem cells are potential sources for cell transplantation into patients to treat serious degenerative conditions. For example, Parkinson’s disease results from a progressive loss of cells that produce the neurotransmitter dopamine in a specific region of the brain. Transplantation of stem cells that have been induced to differentiate as dopamine-producing cells holds great promise as an effective long-term treatment. Similarly, stem cells may become a source of insulin-producing cells for transplantation into the pancreas of individuals with diabetes mellitus. Stem cells might also provide replacement neurons in people with spinal cord injury or other types of neurological damage. Researchers ultimately hope to establish lines of human pluripotent stem cells that can grow indefinitely in culture, be induced to differentiate under controlled conditions and stably maintain their differentiated state, and be manipulated genetically. They particularly want to develop ES cell lines from patients with cancer, diabetes, cardiovascular disease, and neurogenerative disorders (such as Parkinson’s disease); such cell lines would be invaluable in research on these disorders. Although work on stem cells in mice and other mammals has been conducted for many years, similar studies in humans have progressed slowly, despite the great promise of stem cells as a therapeutic tool. Thus far, the only known source of ES cells is early human embryos left over from in vitro fertilization (see Chapter 50). Some countries have government restrictions on public funding due to ethical considerations related to the origins of ES cells. Other countries, notably some Asian countries, have no restrictions on this type of research. In the United States, the Obama administration lifted restrictions on stem cell research that were established during the George W. Bush administration, and public funding became available in 2009.

Volker Steger/Science Photo Library/Photo Researchers, Inc.

Smooth muscle cells

Blastocyst 1

Human embryonic stem (ES) cells are derived from 5- to 6-day-old embryo (blastocyst).

2 ES cells are present in liquid drops of this stem

Neuron

Red blood cells 3

cell culture.

It may be possible to induce ES cells to differentiate into any of approximately 250 cell types of the human body.

FIGURE 17-5 Human embryonic stem (ES) cells

Induced pluripotent stem (iPS) cells may eventually replace ES cells In 2007 and 2008, scientists at several research labs were able to produce induced pluripotent stem (iPS) cells by introducing a few key transcription factors into mature mouse and human cells such as fibroblasts and skin cells. (Recall from Chapter 14 that transcription factors are DNA-binding proteins that regulate gene transcription in eukaryotic cells.) The transcription factors reprogrammed the developmental state of the mature cells so they became pluripotent embryonic cells, but the mechanism by which they accomplished this is currently unknown. The transcription factors may have removed methyl groups from key genes that are normally turned off in adult cells but turned on in embryonic cells. (Recall from Chapter 14 that epigenetic inheritance consists of gene inactivation, for example, by DNA methylation.) For genes that are inactivated by methylation, removing the methyl groups would reactivate the genes. Scientists must learn more about iPS cells before they can be used to replace embryonic stem cells in research. It does not appear that there is any difference between iPS cells and ES cells. If this is the case, then the ethical concerns that make ES cells controversial (discussed in the next section) may soon not matter because scientists will not need to use ES cells.

by a process other than fertilization into a woman’s body. Many countries are opposed to human reproductive cloning. In contrast, human therapeutic cloning would involve duplication of human ES cells or iPS cells for scientific study or medical purposes; no newborn human would develop. These cells would provide a supply of replacement tissues, such as heart muscle or nervous tissue, for revolutionary medical procedures. Many people support research in therapeutic cloning because of the potential benefits in curing disease. Other people have ethical or religious objections if ES cells are used.

Review ■ ■

■

What lines of evidence support the principle of nuclear equivalence? Why was Wilmut’s team successful in mammalian cloning where others had failed? What are stem cells?

17.2 THE GENETIC CONTROL OF DEVELOPMENT ■ ■ LEARNING OBJECTIVES 4 Indicate the features of Drosophila melanogaster, Caenorhabditis elegans,

Ethical questions exist relating to human cloning Cloning research continues to fuel an ongoing debate regarding the potential for human cloning and its ethical and religious implications. In the United States, the National Bioethics Advisory Commission has been established to study this and other questions. In considering these issues, it is important to recognize that cloning is a broad term that includes several different processes involved in producing biological cells, tissues, organs, or organisms. Human reproductive cloning has the goal of producing a newborn human that is genetically identical to another, usually adult, human. It would involve placing a human embryo produced

5

Mus musculus, and Arabidopsis thaliana that have made these organisms valuable models in developmental genetics. Distinguish among maternal effect genes, segmentation genes, and homeotic genes in Drosophila.

6 Explain the relationship between transcription factors and genes that 7

control development. Define induction and apoptosis, and give examples of the roles they play in development.

Development has been an important area of research for many years, and biologists have spent considerable time studying the development of invertebrate and vertebrate animals. By investigating

patterns of morphogenesis in different species, researchers have identified both similarities and differences in the basic plan of development from a zygote to an adult in organisms ranging from the sea urchin to mammals (see Chapter 51). In addition to descriptive studies, many classic experiments have demonstrated how groups of cells differentiate and undergo pattern formation. Researchers have developed elaborate screening programs to detect mutations that let them identify many developmental genes in both plants and animals. They then use molecular genetic techniques and other sophisticated methodologies to determine how those genes work and how they interact to coordinate developmental processes.

A variety of model organisms provide insights into basic biological processes In studies of the genetic control of development, the choice of organism to use as an experimental model is important. A model organism is a species chosen for biological studies because it has characteristics that allow for the efficient analysis of biological processes. Because most model organisms are small organisms with short generation times, they are easy to grow and study under controlled conditions. For example, mice are a better model organism than kangaroos. One of the most powerful approaches in developmental genetics involves isolating mutants of a model organism with abnormal development. Not all organisms have useful characteristics that allow researchers to isolate and maintain developmental mutants for future study. Geneticists so thoroughly understand the genetics of the fruit fly, Drosophila melanogaster, that this organism has become one of the most important systems for such studies. Other organisms—the yeast Saccharomyces cerevisiae; the nematode worm, Caenorhabditis elegans; the zebrafish, Danio rerio; the laboratory mouse, Mus musculus; and certain plants, including Arabidopsis thaliana, a tiny weed—have also become important models in developmental genetics. Each of these organisms has attributes that make it particularly useful for examining certain aspects of development (FIG. 17-6). In the 1990s, developmental geneticists working on C. elegans discovered RNA interference, which has become a powerful research tool. In RNA interference (RNAi) certain small RNA molecules interfere with the expression of genes or their RNA transcripts (see Chapter 13). One way these molecules work is to silence genes by selectively cleaving mRNA molecules that have base sequences complementary to the small RNA molecules. The use of RNAi makes it possible to knock out the expression of a specific gene in an organism during its development, thereby deducing the purpose of the gene.

Many genes that control development have been identified in the fruit fly

The Drosophila life cycle includes egg, larval, pupal, and adult stages Development in Drosophila consists of several distinct stages (FIG. 17-7). After the egg is fertilized, a period of embryogenesis occurs during which the zygote develops into a sexually immature form known as a larva (pl., larvae). After hatching from the egg, each larva undergoes several molts (shedding of the external covering, or cuticle). The periods between molts are called instars. Each molt results in a size increase until the larva is ready to become a pupa. Pupation involves a molt and the hardening of the new external cuticle so that the pupa is completely encased. The insect then undergoes metamorphosis, a complete change in form. During that time, most of the larval tissues degenerate and other tissues differentiate to form the body parts of the sexually mature adult fly. The larvae are wormlike in appearance and look nothing like the adult flies. However, very early in embryogenesis of the developing larvae, precursor cells of many of the adult structures are organized as relatively undifferentiated, paired structures called imaginal discs. The name comes from imago, the adult form of the insect. Each imaginal disc occupies a definite position in the larva and will form a specific structure, such as a wing or a leg, in the adult body (FIG. 17-8). The discs are formed by the time embryogenesis is complete and the larva is ready to begin feeding. In some respects the larva is a developmental stage that feeds and nurtures the precursor cells that give rise to the adult fly, which is the only form that reproduces. The organization of the precursors of the adult structures, including the imaginal discs, is under genetic control. Thus far, more than 50 genes have been identified that specify the formation of the imaginal discs, their positions within the larva, and their ultimate functions within the adult fly. Those genes were identified through mutations that either prevent certain discs from forming or alter their structure or ultimate fate. Drosophila developmental mutations affect the body plan Scientists have identified many developmental mutations in Drosophila. Researchers have examined their effects on development in various combinations and have studied them extensively at the molecular level. In our discussion we pay particular attention to mutations that affect the segmented body plan of the organism, in both the larva and the adult.

▲

The fruit fly (Drosophila) genome sequence, which was completed in late 1999, includes about 13,600 protein-coding genes. At least 1200 of these are essential for embryonic development. Two of the traditional advantages of Drosophila as a research organism are the abundance of mutant alleles, including those of

developmental genes, available for study and the relative ease with which a new mutation is mapped on the chromosomes. U.S. biologist Edward B. Lewis (1918–2004), a pioneering developmental geneticist, began working with Drosophila mutants in the 1940s. The work of German researcher Christiane Nüsslein-Volhard and American Eric Wieschaus extended our understanding of development in fruit flies. Lewis, Nüsslein-Volhard, and Wieschaus shared the 1995 Nobel Prize in Physiology or Medicine for their decades of painstaking research on the genetics of Drosophila development. Many of the genes they discovered in the fruit fly are now known to be important in the growth and development of all animals.

FIGURE 17-6 Model organisms in developmental genetics

Herman Eisenbeiss/Photo Researchers, Inc.

Microfield Scientific Ltd./Science Photo Library/Photo Researchers, Inc.

5 μm

(b) Fruit fly (Drosophila melanogaster) development involves an anterior-posterior body plan that is common to many invertebrates and vertebrates.

Mark Smith/Photo Researchers, Inc.

Sinclair Stammers/Science Photo Library/Photo Researchers, Inc.

(a) Yeast (Saccharomyces cerevisiae) is a model organism for studying developmental genetics in haploid, unicellular eukaryotes.

0.25 mm

Dr. Jeremy Burgess/Photo Researchers, Inc.

(d) Zebrafish (Danio rerio) are small (2 to 4 cm long) and easy to grow, providing an ideal organism for studying the genetic basis of development in vertebrates.

J. M. Labat/Photo Researchers, Inc.

(c) The nematode (Caenorhabditis elegans) adult body consists of only 959 cells, and its transparency allows researchers to trace the development of each cell.

(e) Mouse (Mus musculus) is a model organism for studying developmental genetics, including the genetics of cancer, in mammals.

(f) Mouse-ear cress (Arabidopsis thaliana) is a small plant with a small genome that is used to study development in flowering plants.

Adult

Male

Female Sperm

Labium Mouthparts Dorsal Eye Second Wing Haltere Genital prothorax leg apparatus

Egg Adult 31/2 to 41/2 days

Fertilized egg 1 day

Pupa

Larva

FIGURE 17-8 The location of imaginal discs Each pair of discs in a Drosophila larva (bottom) develops into a specific pair of structures in the adult fly.

21/2 to 3 days 1 day Third instar larva

1 day

First instar larva

Second instar larva

FIGURE 17-7 The life cycle of Drosophila As it develops from a zygote to a sexually mature adult fly, a fruit fly passes through several stages. It takes about 12 days, at 25°C, to complete the life cycle. The dotted lines within the pupa represent the animal undergoing metamorphosis.

In early development of Drosophila, the structure of the egg becomes organized as it develops in the ovary of the female. Stores of mRNA, along with yolk proteins and other cytoplasmic molecules, pass from the surrounding maternal cells into the egg. Immediately after fertilization, the zygote nucleus divides, beginning a series of 13 mitotic divisions. Each division takes only 5 or 10 minutes, which means that the DNA in the nuclei replicates constantly and at a very rapid rate. During that time, the nuclei do not synthesize RNA. Cytokinesis does not take place, and the nuclei produced by the first seven divisions remain at the center of the embryo until the eighth division occurs. At that time, most of the nuclei migrate out from the center and become localized at the periphery of the embryo. This is known as the syncytial blastoderm stage because the nuclei are not surrounded by individual plasma membranes. (A syncytium is a structure containing many nuclei residing in a common cytoplasm.) Subsequently, plasma membranes form, and the embryo becomes known as a cellular blastoderm. Maternal Effect Genes The genes that organize the structure of the egg cell are called maternal effect genes. These genes in the surrounding maternal tissues are transcribed to produce mRNA molecules that are transported into the developing egg. Analysis of

mutant flies with defective maternal effect genes has revealed that many of these genes are involved in establishing the polarity of the embryo—such as which part of the embryo will become the head. Polarity dictates those parts of the egg that are dorsal or ventral and those that are anterior or posterior (see Chapter 30); thus, these genes are known as egg polarity genes. FIGURE 17-9a illustrates concentration gradients for two specific maternal mRNA molecules in the very early embryo. These mRNA transcripts of some of the maternal effect genes are identified by their ability to hybridize with radioactive DNA probes derived from cloned genes (discussed in Chapter 15). Alternatively, researchers use fluorescently tagged antibodies (as in the chapter opening photograph) to bind to specific protein products of the maternal effect genes. These protein gradients organize the early pattern of development in the embryo by determining anterior and posterior regions. A combination of protein gradients may provide positional information that specifies the fate—that is, the developmental path—of each nucleus within the embryo. For example, mutations in certain maternal effect genes result in an embryo with two heads or two posterior ends. In many cases, injecting normal maternal mRNA into the mutant embryo reverses the phenotype associated with a mutation in a given maternal effect gene. The fly subsequently develops normally, indicating that the gene product is needed only for a short time in the earliest stages of development. Segmentation Genes As the nuclei start to migrate to the periph-

ery of the embryo, segmentation genes in those nuclei begin to produce embryonic mRNA. Thus far, geneticists have identified at least 24 segmentation genes that are responsible for generating a repeating pattern of body segments within the embryo and adult fly. Based on the study of mutant phenotypes, researchers group the segmentation genes into three classes—gap genes, pair–rule genes, and segment polarity genes. Gap genes are the first set of segmentation genes to act. These genes interpret the maternal anterior-posterior information in the

Stage of development Anterior

Posterior

Gene activity Anterior

Posterior

+ ++ + + + + ++

(a) At 1.25 hours after fertilization, the embryo consists of a common cytoplasm with about 128 nuclei (purple circles). Maternal effect genes divide the embryo into anterior and posterior sections. The crosses in the anterior region represent mRNA molecules transcribed from one maternal effect gene. The pink shading represents a different maternal mRNA molecule that is more concentrated in the anterior region.

+ + +

++

FIGURE 17-9 Early development in Drosophila Longitudinal sections on the left show two early stages of development of the Drosophila embryo. These are matched with the simplified patterns of gene activity at each stage on the right. (Adapted from Akam, M. E., "The Molecular Basis for Metameric Pattern in the Drosophila Embryo," Development, Vol. 101, 1987.)

egg and begin organization of the body into anterior, middle, and posterior regions (FIG. 17-9b). A mutation in one of the gap genes usually causes the absence of one or more body segments in an embryo (FIG. 17-10a). The other two classes of segmentation genes do not act on small groups of body segments; instead, they affect all segments. Mutations in pair–rule genes delete every other segment, producing a larva with half the normal number of segments (FIG. 17-10b

(a) Gap genes

Wild-type

(b) Pair–rule genes

Mutant Wild-type

(c) Segment polarity genes

Gap genes, pair–rule genes, and segment polarity genes control the pattern of body segments in a Drosophila embryo. The blue bands mark the regions in which the protein products of these genes are normally expressed in wild-type embryos. These same regions are absent in embryos in which the gene is mutated—that is, the mutant lacks the blue-shaded parts. The resulting phenotype is characteristic of the class to which the gene belongs. (Adapted from Nusslein-Volhard, C., and E. Wieschaus, "Mutations Affecting Segment Number and Polarity in Drosophila," Nature, Vol. 287, 1980.)

and FIG. 17-11). Mutations in segment polarity genes produce segments in which one part is missing and the remaining part is duplicated as a mirror image (FIG. 17-10c). The effects of the different classes of mutants are summarized in TABLE 17-1. Each segmentation gene has distinctive times and places in the embryo in which it is most active. The observed pattern of expression of maternal effect genes and segmentation genes indicates that a progressive series of developmental events determines cells destined to form adult structures. First, maternal effect genes that form gradients of morphogens in the egg determine the anteriorposterior (head-to-tail) axis and the dorsal and ventral regions of the embryo. A morphogen is a chemical agent that affects the differentiation of cells and the development of form. Morphogen gradients are signals that help cells determine their location within the embryo and promote their eventual differentiation into specialized tissues and organs.

Classes of Genes Involved in Establishing Segmented Body Plan in Drosophila

Class of Gene Maternal effect genes

Segmentation genes Gap genes Pair–rule genes Segment polarity genes Homeotic genes

Wild-type

Mutant

FIGURE 17-10 A comparison of mutations in Drosophila segmentation genes

++

(b) At 2 hours after fertilization, about 1500 nuclei have migrated into the periphery of the embryo and started to make their own mRNA molecules. The gap genes divide the embryo into anterior, middle, and posterior sections. The maternal RNA (shown in pink) is now transcribed from segmentation genes (crosses) in nuclei in the anterior region. The mRNA from another gap gene is transcribed in the middle of the embryo (black region).

TABLE 17-1

Mutant

Site of Gene Activity

Functions of Genes

Maternal tissues surrounding egg

Initiate anterior-posterior regions in embryo and regulate expression of segmentation genes

Embryo Embryo Embryo

Divide embryo into broad regions and influence activity of pair–rule genes Divide embryo into strips, each about two segments wide, and influence activity of segment polarity genes Define all segments of body, and divide each segment into anterior and posterior halves

Embryo

Control identities of segments; expression of homeotic gene is controlled by segmentation genes

Steven W. Paddock

50 μm

FIGURE 17-11 Segmentation gene activity The bright bands in this fluorescence LM reveal the presence of mRNA transcribed from one of the pair–rule genes known as fushi tarazu (Japanese for “not enough segments”). When this locus is mutated, the segments of the larva that are normally derived from these bands are absent.

Next, segmentation genes respond to the morphogens at each location to regulate the production of segments from the head to the posterior region. Within each segment, other genes are then activated to specify which body part that segment will become. Every cell’s position is further specified with a specific “address” designated by combinations of the activities of the regulatory genes. The segmentation genes act in sequence, with the gap genes acting first, then the pair–rule genes, and finally the segment polarity genes. In addition, the genes of the three groups interact. Each time a new group of genes acts, cells for that group become more determined in their development. As the embryo develops, each region is progressively subdivided into smaller regions.

Antenna

Most segmentation genes code for transcription factors. For example, some segmentation genes code for a “zinc-finger” type of DNA-binding regulatory protein. (See Figure 14-12b to review zinc fingers.) Homeotic genes, discussed next, also code for transcription factors. The fact that many of the genes involved in controlling developmental processes code for transcription factors indicates that those proteins act as genetic switches to regulate the expression of other genes. Once researchers have identified proteins that function as transcription factors, they can use the purified proteins to determine the DNA target sequences to which the proteins bind. This approach has been increasingly useful in identifying additional parts of the regulatory pathway involved in different stages of development. Transcription factors also play a role in cancer (discussed later in the chapter). Homeotic Genes After segmentation genes have established the basic pattern of segments in the fly body, homeotic genes specify the developmental plan for each segment. Mutations in homeotic genes cause one body part to be substituted for another and therefore produce some peculiar changes in the adult. A striking example is the Antennapedia mutant fly, which has legs that grow from the head where the antennae would normally be (FIG. 17-12). Homeotic genes in Drosophila were originally identified by the altered phenotypes produced by mutant alleles. When geneticists Eye

Homeotic leg

Homeotic leg

Dr. Thomas Kaufman

Eye

Mouthparts Normal adult fly

Homeotic Antennapedia mutant

(a) The head of a normal fly and a fly with an Antennapedia mutation.

250 μm

(b) SEM of the head of a fly with an Antennapedia mutation.

FIGURE 17-12 The Antennapedia locus Antennapedia mutations cause homeotic transformations in Drosophila in which legs or parts of legs replace the antennae.

analyzed the DNA sequences of several homeotic genes, they discovered a short DNA sequence of approximately 180 base pairs, which characterizes many homeotic genes as well as some other genes that play a role in development. This DNA sequence is called the homeobox. Each homeobox codes for a protein functional region called a homeodomain, consisting of 60 amino acids that form four a-helices. One of these serves as a recognition helix that binds to specific DNA sequences and thereby affects transcription. Thus, the products of homeotic genes, like those of the earlieracting segmentation genes, are transcription factors. In fact, some segmentation genes also contain homeoboxes. Studies of Hox genes, clusters of homeobox-containing genes that specify the anterior-posterior axis during development, provide insights about evolutionary relationships. Hox genes were initially discovered in Drosophila, where they are arranged in two adjacent groups on the chromosome: the Antennapedia complex and the bithorax complex. As Hox genes have been identified in other animals, such as other arthropods, sea anemones, annelids (segmented worms), roundworms, and vertebrates, researchers have found that these genes are also clustered and that their organization is remarkably similar to that in Drosophila. FIGURE 17-13 compares the organization of the Hox gene clusters of the fruit fly and mouse. The fruit fly and mouse Hox genes are located in the same order along the chromosome. Moreover, the linear order of the genes on the chromosome reflects the order of the corresponding regions they control, from anterior to posterior, in the animal. This organization apparently reflects the need for these genes to be transcribed in a specific temporal sequence. Drosophila has only one Antennapedia–bithorax complex. However, mice, humans, and many other vertebrates have four similar Hox gene clusters, each located in a different chromosome. These complexes probably arose through gene duplication. The fact that extra copies of these genes are present helps explain why mutations causing homeotic-like transformations are seldom seen in vertebrate animals. One does not see an extra eye growing where a mouse leg should be, for example. However, one particular type of Hox mutation that has been described in both mice and humans causes abnormalities in the limbs and genitalia. The involvement of the genitalia provides a further explanation for the rarity of these mutant alleles because affected individuals are unlikely to reproduce. The fact that very similar developmental controls are seen in organisms as diverse as insects and vertebrates (including humans) indicates that the basic mechanism evolved early and has been highly conserved in all animals that have an anterior-posterior axis, even those that are not segmented. Clearly, once a successful way of regulating groups of genes and integrating their activities evolved, it was retained, although it has apparently been modified to provide alterations of the body plan. The finding of homeobox-like genes in plants suggests these genes originated early during eukaryotic evolution. With further investigations, researchers hope to develop an overall model of how the rudiments of morphogenesis are controlled in all multicellular eukaryotes. These systems of master genes that regulate development are a rich source of “molecular fossils” that are illuminating evolutionary history in new and exciting ways. These kinds of studies have led to an ongoing synthesis of evolution and developmental biology that has come to be known as Evo Devo.

KEY POINT

Hox genes are arranged in the same order on the chromosome as they are expressed along the anterior-posterior axis of the embryo.

Anterior region

Posterior region

Fruit fly adult

Fruit fly embryo (10 hours) Fruit fly chromosome

1 2 3 4 5 6 7

9 10 11

13

1 2 3 4 5 6 7 8 9 Mouse chromosomes

4 5 6 1

3 4

8 9 10 11 12 13 8 9 10 11 12 13

Mouse embryo

Mouse adult

FIGURE 17-13 Hox gene clusters Clusters of Hox genes are found in all animal groups except sponges. Note that in each organism, the linear sequence of these developmental genes (color-coded bands) on the chromosome(s) reflects their spatial order of expression in the embryo.

Caenorhabditis elegans has a relatively rigid developmental pattern The nematode worm C. elegans is an ideal model organism because its system for the genetic control of development is relatively easy to study. Sydney Brenner, a British molecular geneticist, began studying molecular development in this animal at Cambridge

0.3 mm

James King-Holmes/Photo Researchers, Inc.

University in the 1960s. He selected C. elegans because it is small, has a short life cycle (its normal lifespan is about 3 weeks), and is genetically well characterized. Consisting of about 19,700 proteincoding genes, it was the first animal genome sequenced. Today, C. elegans is an important tool for answering basic questions about the development of individual cells within a multicellular organism. When an adult, C. elegans is only 1.5 mm long and contains only 959 somatic cells (FIG. 17-14). Individuals are either males or hermaphrodites, organisms with both sexes in the same individual. Hermaphroditic C. elegans are capable of self-fertilization, which makes it easy to obtain offspring that are homozygous for newly induced recessive mutations. The availability of males that can reproduce sexually with the hermaphrodites makes it possible to perform genetic crosses as well. Because the worm’s body is transparent, researchers can follow the development of literally every one of its somatic cells using a Nomarski differential interference microscope, which provides contrast in transparent specimens. As a result of efforts by several research teams, the lineage of each somatic cell in the adult has been determined. Those studies have shown that the nematode has a very rigid, or fixed, developmental pattern. After fertilization, the egg undergoes repeated divisions, producing about 550 cells that make

(a) False-color scanning optical micrograph of adult hermaphrodite nematodes. The oval structures are eggs. Pharynx

Intestine

Eggs Rectum

Oviduct

Oocytes Uterus Vulva

Anus

(b) Structures in the adult hermaphrodite. The sperm-producing structures are not shown.

FIGURE 17-14 Caenorhabditis elegans This transparent organism has a fixed number of somatic cells. (Part b adapted from Fig. 22-6a in V. Walbot and N. Holder, Developmental Biology. © 1987. Used with permission from McGraw-Hill companies.)

up the small, sexually immature larva. After the larva hatches from the egg case, further cell divisions give rise to the adult worm. The lineage of each somatic cell in the adult can be traced to a single cell in a small group of founder cells, which are formed early in development (FIG. 17-15a). If a particular founder cell is destroyed or removed, the adult structures that would normally develop from that cell are missing. Such a rigid developmental pattern, in which the fates of the cells become restricted early in development, is referred to as mosaic development. Each cell has a specific fate in the embryo, just as each tile in a mosaic design forms part of the pattern. Scientists initially hypothesized that every organ system in C. elegans might be derived from only one founder cell. Detailed analysis of cell lineages, however, reveals that many of the structures found in the adult, such as the nervous system and the musculature, are in fact derived from more than one founder cell (FIG. 17-15b). A few lineages have been identified in which a neuron and a muscle cell are derived from the division of a single cell. A researcher who uses microscopic laser beams small enough to destroy individual cells can determine the influence one cell has on the development of a neighboring cell. Consistent with the rigid pattern of cell lineages, the destruction of an individual cell in C. elegans results, in most cases, in the absence of all of the structures derived from that cell but with the normal differentiation of all the neighboring somatic cells. This observation suggests that development in each cell is regulated through its own internal program. However, the developmental pattern of C. elegans is not entirely mosaic. In some cases, cell differentiation is influenced by interactions with particular neighboring cells, a phenomenon known as induction. One example is the formation of the vulva (pl., vulvae), the reproductive structure through which the eggs are laid. A single nondividing cell, called the anchor cell, is a part of the ovary, the structure in which the germ line cells undergo meiosis to produce the eggs. The anchor cell attaches to the ovary and to a point on the outer surface of the animal, triggering the formation of a passage through which the eggs pass to the outside. When the anchor cell is present, it induces cells on the surface to form the vulva and its opening. If the anchor cell is destroyed by a laser beam, however, the vulva does not form, and the cells that would normally form the vulva remain as surface cells (FIG. 17-16). The analysis of worm mutations has contributed to our understanding of inductive interactions. For example, several types of mutations cause more than one vulva to form. In such mutant animals, multiple vulvae form even if the anchor cell is destroyed. Thus, the mutant cells do not require an inductive signal from an anchor cell to form a vulva. Evidently, in these mutants the gene or genes responsible for vulva formation are constitutive. Conversely, mutants lacking a vulva are also known. In some of these, the cells that would normally form the vulva apparently do not respond to the inducing signal from the anchor cell. C. elegans is a model system to study apoptosis During normal development in C. elegans, there are instances in which cells are destined to die shortly after they are produced. Apoptosis, or genetically programmed cell death, has been observed in a wide variety of organisms, both plant and animal. For example, the human hand is formed as a webbed structure, but

0

The nematode worm C. elegans has a fixed developmental pattern in which the fate of each adult cell can be traced to one of several founder cells in the embryo. 0

Egg

50

100

(a) All somatic cells of C. elegans are derived from five somatic founder cells (shown in blue) produced during the early cell divisions of the embryo. The cell shown in white gives rise to germ line cells.

Time after fertilization (hours)

Time after fertilization (min)

KEY POINT

EGG

Nervous system Hypodermis Musculature

Musculature Nervous system Gonad

Germ line

Hypodermis Nervous system Musculature

10 Hatching Intestine ANTERIOR

POSTERIOR

Intestine

(b) The nervous system, hypodermis, and musculature derive from several founder cells. This lineage map traces the development of the cells that form the intestine. (The dashed lines represent many cell divisions of a particular lineage.)

FIGURE 17-15 Cell lineages of Caenorhabditis elegans

the fingers become individualized when the cells between them undergo apoptosis. In C. elegans, as in other organisms, apoptosis is under genetic control. The worm embryos undergo mitosis to produce a total of 1090 cells, but 131 of these undergo apoptosis during development, resulting in adult worms with 959 cells. In 1986, U.S. molecular geneticist Robert Horvitz discovered several mutant worms that were defective in some way relating to apoptosis. He characterized the mutations and the genes responsible for them at the molecular level. For example, one of the genes codes for a protein that regulates the release from mitochondria of molecules that trigger apoptosis (mitochondria are now known to have a crucial role in apoptosis in animals ranging from C. elegans to humans). Other genes were identified that code for a family of proteins known as caspases, proteolytic enzymes active in the early stages of apoptosis. Homologous genes were subsequently identified in other organisms, including humans; some of these loci in mammals also code for caspases. The molecular events that trigger apoptosis remain an area of intense research interest, the results of which will shed considerable light on the general processes of cell aging and cancer. Robert Horvitz and British scientists Sydney Brenner and John Sulston shared the Nobel Prize in Physiology or Medicine in 2002 for their work on the genetic regulation of organ development and apoptosis in C. elegans. Sulston was one of Brenner’s ▲

FIGURE 17-16 Induction

A single anchor cell induces neighboring cells to form the vulva in C. elegans. This diagram shows how laser destruction of single cells or groups of cells influences neighboring cells.

C. elegans

Gonad

Anchor cell

Anchor cell killed

Developing vulva

(a) Normal development of vulva

All gonadal cells except anchor cell are killed

Developing vulva

(b) No vulva develops

(c) Normal development of vulva

students. He traced the lineage of how a single zygote gives rise to the 959 cells in the nematode adult and also observed that some cells die during normal development. Continuing Sulston’s work on apoptosis, Horvitz, as mentioned earlier, was the first to discover genes involved in apoptosis.

The mouse is a model for mammalian development Mammalian embryos develop in markedly different ways from the embryos of Drosophila and C. elegans. The laboratory mouse, Mus musculus, is the best-studied example of mammalian development. Researchers have identified numerous genes affecting development in the mouse. The mouse genome sequence, which was published in 2002, contains 27,000 to 30,000 genes, similar to the number of protein-coding genes in the human genome. Indeed, 99% of the genes in the mouse have counterparts in humans. The early development of the mouse and other mammals is similar in many ways to human development. During the early developmental period, the embryo lives free in the reproductive tract of the female. It then implants in the wall of the uterus, after which the mother meets its nutritional and respiratory needs. Consequently, mammalian eggs are very small and contain little in the way of food reserves. Almost all research on mouse development has concentrated on the stages leading to implantation because in those stages the embryo is free-living and can be experimentally manipulated. During that period, developmental commitments that have a significant effect on the future organization of the embryo take place. After fertilization, a series of cell divisions gives rise to a loosely packed group of cells. Research shows that all cells are equivalent

in the very early mouse embryo. For example, at the two-cell stage of mouse embryogenesis, if researchers destroy one cell and implant the remaining cell into the uterus of a surrogate mother, a normal mouse usually develops. Conversely, if two embryos at the eight-cell stage of development are fused and implanted into a surrogate mother, a normalsized mouse develops (FIG. 17-17). By using two embryos with different genetic characters, such as coat color, researchers demonstrated that the resulting mouse has four genetic parents. These chimeric mice have fur with patches of different colors derived from clusters of genetically different cells. A chimera is an organism containing two or more kinds of genetically dissimilar cells arising from different zygotes. (The term is derived from the name of a mythical beast that was said to have the head of a lion, the body of a goat, and a snakelike tail.) Chimeras let researchers use genetically marked cells to trace the fates of certain cells during development. The responses of mouse embryos to such manipulations contrast with the mosaic or predetermined nature of early C. elegans development, in which destroying one of the founder cells results in the loss of a significant portion of the embryo. For this reason, biologists say that the mouse has highly regulative development— the early embryo acts as a self-regulating whole that accommodates missing or extra parts. Transgenic mice are used in studies of developmental regulation In transformation experiments, foreign DNA injected into fertilized mouse eggs is incorporated into the chromosomes and expressed (FIG. 17-18). (Also see Figure 15-14 for an example of a transgenic mouse.) The resulting transgenic mice provide insights

8-cell stage white mouse embryo

8-cell stage black mouse embryo

2 1

Enzymes are used to remove thick covering surrounding each embryo.

Embryos are fused.

3

Fused embryo continues to develop.

4

Embryo implanted in uterus of surrogate mother.

FIGURE 17-17 Chimeric mouse Researchers remove embryos from females of two different strains and combine the cells in vitro. The resulting aggregate embryo continues to develop and is implanted in the uterus of a surrogate mother. The offspring has four different genetic parents. Although the surrogate mother is the birth mother, she is not genetically related.

5

Baby mouse has 4 genetic parents and 1 birth parent (surrogate mother).

into how genes are activated during development. Scientists can identify a transgene (foreign gene) that has been introduced into a mouse and determine whether it is active by marking the gene in several ways. Sometimes a similar gene from a different species is used; its protein is distinguished from the mouse protein by specific antibodies. It is also possible to construct a hybrid gene consisting of the regulatory elements of a mouse gene and part of another, nonmouse, gene that codes for a “reporter” protein. For example, the reporter protein could be an enzyme not normally found in

RESEARCH METHOD Why Is It Used?

This method is used to establish a transgenic line of mice that contain genes with useful traits for scientific and medical research.

How Is It Done?

Fertile

1

Collect fertilized eggs.

2

3

Inject cloned DNA fragment into nucleus.

Implant injected embryos into female.

Surrogate mother

4

Test for transgene in offspring of surrogate mother. Transgenic offspring

5

Fetal/neonatal

Pup

Breed transgenic animals.

Adult

FIGURE 17-18 Producing a transgenic mouse

mice. Such studies have been important in showing which DNA sequences of a mouse homeobox gene determine where the gene is expressed in the embryo. Many developmentally controlled genes introduced into mice have yielded important information about gene regulation. When researchers introduce developmentally controlled genes from other species, such as humans or rats, into mice, the genes are regulated the same way they normally are in the donor animal. When introduced into the mouse, for example, human genes encoding insulin, globin, and crystallin—which are normally expressed in cells of the pancreas, blood, and eye lens, respectively—are expressed only in those same tissues in the mouse. That these genes are correctly expressed in their appropriate tissues indicates that the signals for tissue-specific gene expression are highly conserved through evolution. Information on the regulation of genes controlling development in one organism can have valuable applications to other organisms, including humans. Mice and other model organisms are providing insights into the aging process Aging, which is defined as a progressive decline in the performance of various parts of the body, is an important field of developmental biology. The study of aging has great practical potential because age-related diseases represent one of the biggest challenges of biomedical research today. Scientists have demonstrated that many environmental factors influence aging. For example, severe calorie restriction in rodents and other mammals delays aging. However, how the human genome interacts with the cell environment during the aging process is not well understood. Researchers working with model organisms, such as Drosophila and C. elegans, have found that more than 1000 protein-coding genes affect aging to variable degrees. In the nematode worm C. elegans, most aging genes are under the control of three genes (ELT-3, ELT-5, and ELT-6), all of which code for transcription factors. Many of the genes that slow aging in organisms as diverse as yeasts, nematodes, and mammals are associated with cells’ responses to nutrients. Dietary restrictions are known to extend lifespan for many organisms, from yeasts to dogs and possibly even to primates. Two genes (the PHA-4 gene and the SKN-1 gene) that extend survival in C. elegans code for transcription factors. One of the most potent lifespan-extending genes, present in C. elegans, codes for a protein similar to the membrane receptor that allows cells to respond to the peptide insulin-like growth factor (IGF) in humans and other mammals. The binding of IGF to its receptor triggers signal transduction within the cell, which regulates the expression of many genes. Researchers wondered if the IGF receptor influences aging in mammals, including humans. To test this hypothesis, they produced knockout mice with an inactivated gene for the insulin-like growth factor receptor (IGF-1R). (Knockout mice were first discussed in Chapter 15.) Because mice are diploid organisms, they have two copies of the gene, designated igf1r. When both alleles of igf1r are inactivated, the mice develop many abnormalities and die at birth. However, when one allele of igf1r is inactivated and the other is left in its normal state, the mice thrive and live about 25% longer than control mice. In all other respects, the heterozygous mice are normal. They are virtually indistinguishable from control

mice in their rate of development, metabolic rate, ability to reproduce, and body size. Mice heterozygous at the igf1r locus produce fewer IGF receptors, and therefore the cells are not exposed to as much signaling from IGF. Research on the IGF receptor and aging in mice suggests that some of the hundreds of other mutant genes associated with aging in C. elegans may also be involved in mammalian aging. Thus, the genetic control of mammalian aging has enough research topics to occupy molecular geneticists for many years.

Arabidopsis is a model for studying plant development, including transcription factors

Photos courtesy of Jose Luis Reichmann and Elliot Meyerowitz

Mouse-ear cress (Arabidopsis thaliana), the organism most widely used to study the genetic control of development in plants, is a member of the mustard family. Although Arabidopsis itself is a weed of no economic importance, it has several advantages for research. The plant completes its life cycle in just a few weeks and is small enough to be grown in a petri dish, yielding thousands of individuals in limited space. Botanists use chemical mutagens to produce mutant strains and have isolated many developmental mutants. When they insert cloned foreign genes into Arabidopsis cells, the genes integrate into the chromosomes and are expressed. The researchers then induce these transformed cells to differentiate into transgenic plants. In 2000, the Arabidopsis genome became the first plant genome sequenced. Although its genome is relatively small (the rice genome is about four times larger), it includes about 26,000 protein-coding genes. In comparison, Drosophila has about 13,600 genes and C. elegans about 19,700. Many genes in Arabidopsis are functionally equivalent to genes in Drosophila, C. elegans, and other animal species. Of particular importance to development are the more than 1500 genes that code for transcription factors in Arabidopsis (compared to 635 at last count for Drosophila). Not surprisingly, many genes known to specify identities of parts of the Arabidopsis flower

code for transcription factors. During the development of Arabidopsis flowers, four distinct flower parts differentiate: sepals, petals, stamens, and carpels (FIG. 17-19a; also see Fig. 33-1). Sepals cover and protect the flower when it is a bud, petals help attract animal pollinators to the flower, stamens produce pollen grains, and the carpel produces ovules, which develop into seeds following fertilization (discussed in Chapter 28). The ABC model explains how three classes of homeotic genes control the development of the four floral organs; this model may help explain the thousands of different forms of flowers that have evolved. Class A genes are needed to specify sepals; both A and B genes are needed to specify petals; both B and C genes are needed to specify stamens; and class C genes are needed to specify the carpel. Mutations in the A, B, or C organ-identity genes, all of which code for transcription factors, cause one flower part to be substituted for another. For example, class C homeotic mutants (which have an inactive C gene) have petals in place of stamens and sepals in place of the carpel. Therefore, the entire flower consists of only sepals and petals. FIGURES 17-19b, c, and d shows three homeotic mutants of Arabidopsis. The ABC model is not the entire explanation for floral development. Another class of genes, designated SEPALLATA, interacts with the B and C genes to specify the development of petals, stamens, and carpels. Remarkably, when SEPALLATA genes are turned on permanently in Arabidopsis, the resulting plants have white petals growing where the leaves should be. (Because these plants do not photosynthesize, they are grown on a nutrient medium.) These findings in Arabidopsis vastly increase the number of molecular probes available in plants. Researchers use these probes to identify other genes that control development in various plant species and to compare them with genes from a wide range of organisms. The success of the Arabidopsis sequencing project led to an international initiative by plant biologists to understand the functions of all genes in Arabidopsis. This functional genomic information will lead to a far deeper understanding of plant development and evolutionary history.

(a) A normal Arabidopsis flower has four outer leafy green sepals, four white petals, six stamens (the male reproductive structures), and a central carpel (the female reproductive structure).

(b) This homeotic mutant is missing petals.

FIGURE 17-19 Homeotic mutants in Arabidopsis flowers

(c) This homeotic mutant has only sepals and carpels.

(d) This homeotic mutant has sepals and petals but no other floral structures.

Review ■

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What are the relative merits of Drosophila, C. elegans, M. musculus, and Arabidopsis as model organisms for the study of development? What are transcription factors, and how do they influence development? What role does induction play in development? What is apoptosis?

17.3 CANCER AND CELL DEVELOPMENT ■ ■ LEARNING OBJECTIVE 8

Discuss the relationship between cancer and mutations that affect cell developmental processes.

Cancer cells lack normal biological inhibitions. Normal cells are tightly regulated by control mechanisms that prompt them to divide when necessary and prevent them from growing and dividing inappropriately. Cells of many tissues in the adult are normally prevented from dividing; they reproduce only to replace a neighboring cell that has died or become damaged. Cancer cells have escaped such controls and can divide continuously. As a consequence of their abnormal growth pattern, some cancer cells eventually form a mass of tissue called a tumor, or neoplasm. If the tumor remains at the spot where it originated, it can usually be removed by surgery. One of the major problems with certain forms of cancer is that the cells can escape from the controls that maintain them in one location. Metastasis is the spreading of cancer cells to different parts of the body. Cancer cells invade other tissues and form multiple tumors. Lung cancer, for example, is particularly deadly because its cells are highly metastatic; they enter the bloodstream, spread, and form tumors in other parts of the lungs and in other organs, such as the liver and the brain. Biologists now know that cancer is caused by the altered expression of specific genes critical for cell division. Using recombinant DNA methods, researchers have identified many of the genes that when functioning abnormally transform normal cells into cancer cells. The traits of each kind of cancer cell come from at least one, and probably several, oncogenes, or cancer-causing genes. Oncogenes arise from changes in the expression of certain genes called proto-oncogenes, which are normal genes found in all cells and are involved in the control of growth and development. Investigators first discovered oncogenes in viruses that infect mammalian cells and transform them into cancer cells, in a process known as malignant transformation. Such viruses have oncogenes as part of their genomes. When these viruses infect a cell, the viral oncogenes are expressed, causing the cell to divide. A proto-oncogene in a cell that has not been infected by a virus can also mutate and become an oncogene. One of the first oncogenes researchers identified was isolated from a tumor in a human urinary bladder. In the cell that gave rise to the tumor, a protooncogene had undergone a single base-pair mutation; the result was that the amino acid valine replaced the amino acid glycine in the protein product of the gene. This subtle change was apparently a critical factor in converting the normal cell into a cancer cell.

Some of the control mechanisms of cell growth are illustrated in greatly simplified form in FIGURE 17-20. One or more external signal molecules trigger the growth and division of cells. Some of these substances are growth factors that bind to specific growth factor receptors associated with the cell surface, initiating a cascade of events inside the cell. Often the growth factor receptor complex acts as a protein kinase, an enzyme that phosphorylates proteins, which then phosphorylate specific amino acids of several cytoplasmic proteins. This posttranslational modification usually results in the activation of previously inactive enzymes. These activated enzymes then catalyze the activation of certain nuclear proteins, many of which are transcription factors. Activated transcription factors bind to their DNA targets and stimulate the transcription of specific sets of genes that initiate growth and cell division. Even in the simplified scenario presented in the figure, it is evident that multiple steps are required to control cell proliferation. Researchers have identified the proto-oncogenes that encode the products responsible for many of these steps. The current list

KEY POINT

Growth factors are signaling proteins that stimulate cell division in specific target cells.

Growth factor attaches to receptor.

1

Growth factor receptor

Plasma membrane

Cytoplasmic proteins (inactive)

Cytosol

Proteins are phosphorylated (activated).

2

Nuclear envelope 5

Cell growth and division occur.

Nuclear proteins (inactive) 3

4

Nuclear proteins are activated (e.g., activated transcription factors).

Transcription of specific genes is stimulated. Nucleus

FIGURE 17-20 A cell growth control cascade In this example, a growth factor stimulates cell growth. Protooncogenes code for the growth factor receptor and some of the other components of the system. When a proto-oncogene mutates, becoming an oncogene, the cell grows and divides even in the absence of the growth factor.

of known proto-oncogenes includes genes that code for various growth factors or growth factor receptors and genes that respond to stimulation by growth factors, including many transcription factors. When a proto-oncogene mutates or is expressed inappropriately (i.e., when it becomes an oncogene), the cell may misinterpret the signal and respond by growing and dividing. For example, in some cases a proto-oncogene encoding a growth factor receptor mutates in a way that compromises regulation of the receptor. It is always switched “on,” even in the absence of the growth factor that normally controls it. Not all genes that cause cancer when mutated are protooncogenes. About half of all cancers are caused by a mutation in a tumor suppressor gene. These genes, also known as antioncogenes, normally interact with growth-inhibiting factors to block cell division. When mutated, they lose their ability to “put on the brakes,” and uncontrolled growth ensues. Currently, more than 100 oncogenes and 15 tumor suppressor genes have been identified, and many more probably remain to be identified. A change in a single proto-oncogene is usually insufficient to cause a cell to become malignant. The development of cancer is usually a multistep process involving both mutations that activate oncogenes and mutations that inactivate tumor suppressor genes. Additional factors, such as chromosome translocations (see Chapter 16) or the inappropriate activation of the enzyme respon-

■ ■

sible for the maintenance of telomeres (see Chapter 12), may also play a role in the development of cancer. The Cancer Genome Project, a long-term international initiative based in Great Britain, is currently examining every human gene for cancer-related mutations. As more of these genes are discovered and their interactions are unraveled, we will gain a fuller understanding of the control of growth and development. This understanding will improve the diagnosis and treatments of various types of cancer. Some researchers view cancer as a stem cell disease because cancer cells are propagated by a small number of cells that have many properties of stem cells. However, the propagation of normal stem cells is tightly regulated, so the question that some biologists are pursuing is how cell mechanisms that regulate stem cell selfrenewal are changed to cause cancer cell proliferation. It is possible that the relatively long life of stem cells allows them to accumulate mutations that cause the cells to proliferate in an uncontrolled fashion. Although this line of research is promising, it should be emphasized that the validity of cancer as a stem cell disease remains to be demonstrated for most kinds of cancer.

Review How are oncogenes and tumor suppressor genes related to genes involved in the control of normal growth and development? How is a growth factor involved in a typical growth control cascade?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

17.1 (page 370) 1 Distinguish between cell determination and cell differentiation, and between nuclear equivalence and totipotency. An organism contains many types of cells that are specialized to carry out specific functions. These cells are the product of a process of gradual commitment, called cell determination, which ultimately leads to the final step in cell specialization, called cell differentiation. Nuclear equivalence is the concept that, with a few exceptions, ■ all the nuclei of the differentiated somatic cells of an organism are identical to one another and to the nucleus of the single cell from which they descended. Totipotency is the capability of cells to direct the development of an entire organism. 2 Describe the classic experiments of Steward, Gurdon, and Wilmut. ■ Steward induced mature carrot root cells to de-differentiate and express their totipotency, forming an entire plant. ■

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Wilmut fused a differentiated cell from an adult sheep cell with an enucleated egg and implanted it into the uterus of a host mother, where it developed into a normal lamb. Learn more about sheep cloning by clicking on the figure in CengageNOW.

3 Define stem cells, distinguish between embryonic stem cells and pluripotent stem cells, and describe some of the promising areas of research involving stem cells. ■ Stem cells can divide to produce differentiated descendants yet retain the ability to divide to maintain the stem cell population. Totipotent stem cells give rise to all cell types of the body and placenta, whereas pluripotent stem cells give rise to many, but not all, types of cells in an organism. Embryonic stem (ES) cells, formed after a zygote has undergone ■ several rounds of cell division to become a 5- or 6-day-old blastocyst, are pluripotent and have the potential to develop into any type of cell in the body; ES cells are not totipotent because they cannot form cells of the placenta. ■ Stem cells show promise in treating diseases, such as Parkinson’s disease and diabetes mellitus.

17.2 (page 375) 4 Indicate the features of Drosophila melanogaster, Caenorhabditis elegans,

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Gurdon injected nuclei from tadpole intestinal cells into enucleated eggs; a small fraction of these developed into tadpoles, demonstrating the totipotency of the injected nucleus.

Mus musculus, and Arabidopsis thaliana that have made these organisms valuable models in developmental genetics. ■ Developmental mutations have been identified in the fruit fly, Drosophila, many of which affect the organism’s segmented body plan. Many developmental genes discovered in the fruit fly are

now known to be important in the growth and development of all animals.

C. elegans is a roundworm with mosaic development, a rigid developmental pattern in which the fates of cells are restricted early in development. The lineage of every somatic cell in the adult is known, and each can be traced to a single founder cell in the early embryo. ■ The laboratory mouse, M. musculus, is extensively used in studies of mammalian development. The mouse shows regulative development; the very early embryo is a self-regulating whole and can develop normally even if it has extra or missing cells. Transgenic mice, in which foreign genes have been incorporated, have helped researchers determine how genes are activated and regulated during development. ■ Genes affecting development have also been identified in certain plants, such as A. thaliana. The ABC model of interactions among three kinds of genes hypothesizes how floral organs develop in Arabidopsis. Mutations in these homeotic genes cause one flower part to be substituted for another. 5 Distinguish among maternal effect genes, segmentation genes, and homeotic genes in Drosophila. ■ The earliest developmental events to operate in the egg are established by maternal effect genes in the surrounding maternal tissues, which are active prior to fertilization. Some produce gradients of morphogens, chemical agents that affect the differentiation and development of form. Maternal effect genes establish polarity in the embryo. ■ Segmentation genes generate a repeating pattern of body segments within the embryo. Gap genes are segmentation genes that begin organizing the body into anterior, middle, and posterior regions; pair–rule genes and segment polarity genes affect all segments. ■

The later-acting homeotic genes are responsible for specifying the identity of each segment. 6 Explain the relationship between transcription factors and genes that control development. ■ Transcription factors are DNA-binding proteins that regulate transcription in eukaryotes. Some genes that code for transcription factors contain a DNA sequence called a homeobox, which codes for a protein with a DNA-binding region called a homeodomain. ■ Some homeobox genes are organized into complexes that appear to be systems of master genes specifying an organism’s body plan. Parallels exist between the homeobox complexes of Drosophila and those of other animals. 7 Define induction and apoptosis, and give examples of the roles they play in development. ■ Induction is developmental interactions with neighboring cells. During development in C. elegans, the anchor cell induces cells on the surface to organize to form the vulva, the structure through which the eggs are laid. ■ Apoptosis is programmed cell death. During human development, the human hand forms as a webbed structure, but the fingers become individualized when the cells between them die. ■

17.3 (page 387) 8 Discuss the relationship between cancer and mutations that affect cell developmental processes. ■ The traits of cancer cells are due to cancer-causing oncogenes. Oncogenes arise from changes in the expression of normal genes called proto-oncogenes, which exist in all cells and are involved in the control of growth and development. ■ Known proto-oncogenes include genes that code for various growth factors or growth factor receptors and genes that respond to stimulation by growth factors, including many transcription factors. When a proto-oncogene is expressed inappropriately (i.e., becomes an oncogene), the cell may misinterpret the signal and respond by growing and dividing. ■ Tumor suppressor genes normally interact with growth-inhibiting factors to block cell division. A mutation in a tumor suppressor gene may inactivate it and thereby lead to cancer.

T E S T YO U R U N D E R S TA N D I N G 1. Morphogenesis occurs through the multistep process of (a) differentiation (b) determination (c) pattern formation (d) totipotency (e) selection 2. The cloning experiments carried out on frogs demonstrated that (a) all differentiated frog cells are totipotent (b) some differentiated frog cells are totipotent (c) all nuclei from differentiated frog cells are totipotent (d) some nuclei from differentiated frog cells are totipotent (e) cell differentiation always requires the loss of certain genes 3. The anterior-posterior axis of a Drosophila embryo is first established by certain (a) homeotic genes (b) maternal effect genes (c) segmentation genes (d) proto-oncogenes (e) pair–rule genes 4. You discover a new Drosophila mutant in which mouth parts are located where the antennae are normally found. You predict that the mutated gene is most likely a (a) homeotic gene (b) gap gene

(c) pair–rule gene (d) maternal effect gene (e) segment polarity gene 5. Most segmentation genes code for (a) transfer RNAs (b) enzymes (c) transcription factors (d) histones (e) transport proteins 6. Homeobox genes (a) are found in fruit flies but no other animals (b) tend to be expressed in the order that they appear on a chromosome (c) contain a characteristic DNA sequence (d) b and c (e) a, b, and c 7. The developmental pattern of C. elegans is said to be mosaic because (a) development is controlled by gradients of morphogens (b) part of the embryo fails to develop if a founder cell is destroyed (c) some individuals are self-fertilizing hermaphrodites (d) all development is controlled by maternal effect genes (e) apoptosis never occurs

8. Which of the following illustrates the regulative nature of early mouse development? (a) the mouse embryo is free-living prior to implantation in the uterus (b) it is possible to produce a transgenic mouse (c) it is possible to produce a mouse in which a specific gene has been knocked out (d) genes related to Drosophila homeotic genes have been identified in mice (e) a chimeric mouse can be produced by fusing two mouse embryos 9. Arabidopsis is useful as a model organism for the study of plant development because (a) it is of great economic importance (b) it has a very long generation time (c) many developmental mutants have been isolated (d) it contains a large amount of DNA per cell (e) it has a rigid developmental pattern 10. Pluripotent stem cells (a) lose genetic material during development (b) give rise to many, but not all, types of cells in an organism (c) organize into recognizable structures through pattern

formation (d) cannot grow in tissue culture (e) have been used to clone a sheep and several other mammals 11. Which of the following statements about cancer is false? (a) oncogenes arise from mutations in proto-oncogenes (b) tumor suppressor genes normally interact with growth-inhibiting factors to block cell division (c) more than 100 oncogenes and 15 tumor suppressor genes have been identified (d) oncogenes were first discovered in mouse models for cancer (e) the development of cancer is usually a multistep process involving both oncogenes and mutated tumor suppressor genes 12. Proto-oncogenes code for (a) morphogens (b) antibodies for immune responses (c) growth factor receptors and other components of the growth control cascade (d) enzymes such as reverse transcriptase (e) ES cells

CRITICAL THINKING 1. Why is an understanding of gene regulation in eukaryotes crucial to an understanding of developmental processes? 2. Why do scientists study development in more than one type of organism? 3. Could a gene be involved in the growth of both stem cells and some kinds of cancer? Explain your answer.

to specify the carpel. If a mutation occurs in one of the B genes, rendering it inactive, what will the resulting flowers consist of? Sepal A genes Petal B genes Stamen

4. EVOLUTION LINK. How are the striking similarities among genes that govern development in widely differing species strong evidence for evolution? 5. EVOLUTION LINK. What is the common ground between evolutionary biologists and developmental biologists who have adopted the perspective known as Evo Devo? 6. ANALYZING DATA. Flower parts are arranged in four concentric circles: sepals, petals, stamens, and carpel. According to the ABC model of floral organ development in Arabidopsis, class A genes are needed to specify sepals, the A and B genes to specify petals, the B and C genes to specify stamens, and class C genes

Carpel C genes

Additional questions are available in CengageNOW at www.cengage.com/ login.

18

Introduction to Darwinian Evolution

Science Source/Photo Researchers, Inc.

Charles Darwin. This portrait was made shortly after Darwin returned to England from his voyage around the world.

KEY CONCEPTS

A

18.1 Evolution is the accumulation of inherited changes

evolved from a single ancestor during Earth’s long history. Thus, organisms

within populations over time.

that are radically different from one another are in fact distantly related,

18.2 Ideas about evolution originated long before Darwin’s time.

linked through numerous intermediate ancestors to a single, common

18.3 Darwin’s scientific theory of evolution, natural se-

a simple, scientifically testable mechanism to explain the relationship

lection, explained how natural forces in the environment could cause evolution. Natural selection occurs because individuals with traits that make them better adapted to local conditions are more likely to survive and produce offspring than are individuals that are not as well adapted. The modern synthesis combines Darwin’s theory with genetics.

18.4 The evidence that evolution has taken place and is still occurring is overwhelming. This evidence includes fossils, biogeography, comparative anatomy, molecular biology, developmental biology, and evolutionary experiments with living organisms.

great deal of evidence suggests that the biological diversity represented by the millions of species currently living on our planet

ancestor. The British naturalist Charles Darwin (1809–1882) developed among Earth’s diversity of organisms. He argued persuasively that all the species that exist today, as well as the countless extinct species that existed in the past, arose from earlier ones by a process of gradual divergence (splitting into separate evolutionary pathways), or evolution. The concept of evolution is the cornerstone of biology because it links all fields of the life sciences into a unified body of knowledge. As stated by U.S. geneticist Theodosius Dobzhansky, “Nothing in biology makes sense except in the light of evolution.”1 Biologists seek to understand both the remarkable variety and the fundamental similarities of organisms within the context of evolution. The science of evolution allows biologists to 1

American Biology Teacher, Vol. 35, No. 125 (1973).

compare common threads among organisms as seemingly differ-

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ent as bacteria, whales, lilies, slime molds, and tapeworms. Animal behavior, developmental biology, genetics, evolutionary ecology,

20 Percent resistance

systematics, and molecular evolution are examples of some of the biological disciplines that are grounded in evolution. This chapter discusses Charles Darwin and the development of his evolutionary theory by natural selection. It also presents evidence that supports evolution, including fossils, biogeography, comparative anatomy, molecular biology, developmental biology, and experimental studies of ongoing evolutionary change in both

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the laboratory and nature. 0

18.1 WHAT IS EVOLUTION? ■ ■ LEARNING OBJECTIVE 1

Define evolution.

In beginning our study of evolution, we define evolution as the accumulation of genetic changes within populations over time. A population is a group of individuals of one species that live in the same geographic area at the same time. Just as the definition of a gene changed as you studied genetics, you will find that the definition of evolution will become more precise in later chapters. The term evolution does not refer to changes that occur in an individual within its lifetime. Instead, it refers to changes in the characteristics of populations over the course of generations. These changes may be so small that they are difficult to detect or so great that the population differs markedly from its ancestral population. Eventually, two populations may diverge to such a degree that we refer to them as different species. The concept of species is developed extensively in Chapter 20. For now, a simple working definition is that a species is a group of organisms, with similar structure, function, and behavior, that are capable of interbreeding with one another. Evolution has two main perspectives—the minor evolutionary changes of populations usually viewed over a few generations (microevolution, discussed in Chapter 19) and the major evolutionary events usually viewed over a long period, such as formation of different species from common ancestors (macroevolution, discussed in Chapter 20). Evolution has important practical applications. Agriculture must deal with the evolution of pesticide resistance in insects and other pests. Likewise, medicine must respond to the rapid evolutionary potential of disease-causing organisms such as bacteria and viruses (FIG. 18-1). (Significant evolutionary change occurs in a very short time period in insects, bacteria, and other organisms with short lifespans.) Medical researchers use evolutionary principles to predict which flu strains are evolving more quickly, information that scientists need to make the next year’s flu vaccine. Also, researchers developing effective treatment strategies for the human immunodeficiency virus (HIV) must understand its evolution, both within and among hosts. The conservation management of rare and endangered species makes use of the evolutionary principles of population genetics. The rapid evolution of bacteria and fungi in polluted soils is used in the

1990 1992 1994 1996 1998 2000 2002 2004 2006 2007 Year

FIGURE 18-1 Evolution of antibiotic resistance to ciprofloxacin in E. coli These data show an increasing resistance to E. coli in patients with blood and cerebrospinal infections in England, Wales, and Northern Ireland, 1990 to 2007. (Source: Livermore, D. “Zietgeist of Resistance.” The Journal of Antimicrobial Chemotherapy, Vol. 60, i59–i61, 2007. By permission of Oxford University Press.)

field of bioremediation, in which microorganisms are employed to clean up hazardous-waste sites. Evolution even has applications beyond biology. For example, certain computer applications make use of algorithms that mimic natural selection in biological systems.

Review ■ ■

What is evolution? Do individuals evolve? Explain your answer.

18.2 PRE-DARWINIAN IDEAS ABOUT EVOLUTION ■ ■ LEARNING OBJECTIVE 2

Discuss the historical development of evolutionary theory.

Although Darwin is universally associated with evolution, ideas of evolution predate Darwin by centuries. Aristotle (384–322 bce) saw much evidence of natural affinities among organisms. This led him to arrange all the organisms he knew in a “scale of nature” that extended from the exceedingly simple to the most complex. Aristotle visualized organisms as being imperfect but “moving toward a more perfect state.” Some scientific historians have interpreted this idea as the forerunner of evolutionary theory, but Aristotle was vague on the nature of this “movement toward perfection” and certainly did not propose that natural processes drove the process of evolution. Furthermore, modern evolutionary theory now recognizes that evolution does not move toward more “perfect” states or even necessarily toward greater complexity. Long before Darwin’s time, fossils had been discovered embedded in rocks. Some of these corresponded to parts of famil-

iar species, but others were strangely unlike any known species. Fossils were often found in unexpected contexts; for example, marine invertebrates (sea animals without backbones) were sometimes discovered in rocks high on mountains. Leonardo da Vinci (1452–1519) was among the first to correctly interpret these unusual finds as the remains of animals that had existed in previous ages but had become extinct. The French naturalist Jean Baptiste de Lamarck (1744–1829) was the first scientist to propose that organisms undergo change over time as a result of some natural phenomenon rather than divine intervention. According to Lamarck, a changing environment caused an organism to alter its behavior, thereby using some organs or body parts more and others less. Over several generations, a given organ or body part would increase in size if it was used a lot, or shrink and possibly disappear if it was used less. His hypothesis required that organisms pass traits they acquired during their lifetimes to their offspring. For example, Lamarck suggested that the long neck of the giraffe developed when a short-necked ancestor stretched its neck to browse on the leaves of trees. Its offspring inherited the longer neck, which stretched still further as they ate. This process, repeated over many generations, resulted in the long necks of modern giraffes. Lamarck also thought that all organisms were endowed with a vital force that drove them to change toward greater complexity and “perfection” over time. Lamarck’s proposed mechanism of evolution is quite different from the mechanism later proposed by Darwin. However, Lamarck’s hypothesis remained a reasonable explanation for evolution until Mendel’s basis of heredity was rediscovered at the beginning of the 20th century. At that time, Lamarck’s ideas were largely discredited.

Review ■ ■

Why is Aristotle linked to early evolutionary thought? What were Jean Baptiste de Lamarck’s ideas concerning evolution?

18.3 DARWIN AND EVOLUTION ■ ■ LEARNING OBJECTIVES 3 4

Explain the four premises of evolution by natural selection as proposed by Charles Darwin. Compare the modern synthesis with Darwin’s original evolutionary theory.

Darwin, the son of a prominent physician, was sent at the age of 15 to study medicine at the University of Edinburgh. Finding himself unsuited for medicine, he transferred to Cambridge University to study theology. During that time, he became the protégé of the Reverend John Henslow, who was a professor of botany. Henslow encouraged Darwin’s interest in the natural world. Shortly after receiving his degree, Darwin embarked on the HMS Beagle, which was taking a 5-year exploratory cruise around the world to prepare navigation charts for the British navy. The Beagle left Plymouth, England, in 1831 and cruised along the east and west coasts of South America (FIG. 18-2). While other members of the crew mapped the coasts and harbors, Darwin spent many weeks ashore studying the animals, plants, fossils, and geologic formations of both coastal and inland regions, areas that had not been extensively explored. He collected and cataloged thousands of plant and animal specimens and kept notes of his observations, information that became essential in the development of his theory. The Beagle spent almost 2 months at the Galápagos Islands, 965 km (600 mi) west of Ecuador, where Darwin continued his observations and collections. He compared the animals and plants of the Galápagos with those of the South American mainland. He was particularly impressed by their similarities and wondered why the organisms of the Galápagos should resemble those from South America more than those from other islands in different parts of the world. Moreover, although there were similarities between Galápagos and South American species, there were also distinct

Azores Canary Is. Galápagos Is.

Equator Cocos Is. Sydney King George Sound

Cape Verde Is.

Tahiti New Zealand

Hobart

Callao Valparaiso

Ascension Is.

Bahia

St. Helena Is. Mauritius Rio de Janeiro

Montevideo Tierra del Fuego

Falkland Is.

FIGURE 18-2 Animated The voyage of HMS Beagle The 5-year voyage began in Plymouth, England (star), in 1831. Observations made in the Galápagos Islands (bull’s-eye) off the western coast of South America helped Darwin develop a satisfactory mechanism to explain how a population of organisms could change over time.

John Amaldi

differences. There were even recognizable differences in the reptiles and birds from one island to the next. Darwin wondered why these remote islands should have such biological diversity. After he returned home, Darwin pondered these observations and attempted to develop a satisfactory explanation for the distribution of species among the islands. Darwin drew on several lines of evidence when considering how species might have originated. Despite the work of Lamarck, the general notion in the mid-1800s was that Earth was too young for organisms to have changed significantly since they had first appeared. During the early 19th century, however, geologists advanced the idea that mountains, valleys, and other physical features of Earth’s surface did not originate in their present forms. Instead, these features developed slowly over long periods by the geologic processes of volcanic activity, uplift, erosion, and glaciation. On his voyage Darwin took with him Principles of Geology, published by English geologist Charles Lyell in 1830, and studied it carefully. Lyell provided an important concept for Darwin—that the slow pace of geologic processes, which still occur today, indicated that Earth was extremely old. Other important evidence that influenced Darwin was the fact that breeders and farmers could develop many varieties of domesticated animals in just a few generations. They did so by choosing certain traits and breeding only individuals that exhibited the desired traits, a procedure known as artificial selection. Breeders, for example, have produced numerous dog varieties—such as bloodhounds, dalmatians, Airedales, border collies, and Pekinese—by artificial selection. Many plant varieties were also produced by artificial selection. For example, cabbage, broccoli, brussels sprouts, cauliflower, collard greens, kale, and kohlrabi are distinct vegetable crops that are all members of the same species, Brassica oleracea (FIG. 18-3). Selective breeding of the colewort, or wild cabbage, a leafy plant native to Europe and Asia, produced all seven vegetables. Beginning more than 4000 years ago, some farmers artificially selected wild cabbage plants that formed overlapping leaves. Over time, these leaves became so prominent that the plants, which resembled modern cabbages, became recognized as separate and distinct from their wild cabbage ancestor. Other farmers selected different features of the wild cabbage, giving rise to the other modifications. For example, kohlrabi was produced by selection for an enlarged storage stem, and brussels sprouts by selection for enlarged axillary buds. Thus, humans are responsible for the evolution of B. oleracea into seven distinct vegetable crops. Darwin was impressed by the changes induced by artificial selection and hypothesized that a similar selective process occurred in nature. Darwin therefore used artificial selection as a model when he developed the concept of natural selection. The ideas of Thomas Malthus (1766–1834), a British clergyman and economist, were another important influence on Darwin. In An Essay on the Principle of Population as It Affects the Future Improvement of Society, published in 1798, Malthus noted that population growth is not always desirable—a view contrary to the beliefs of his day. He observed that populations have the capacity to increase geometrically (1 ¡ 2 ¡ 4 ¡ 8 ¡ 16) and thus outstrip the food supply, which only has the capacity to increase arithmetically (1 ¡ 2 ¡ 3 ¡ 4 ¡ 5). In the case of humans, Malthus suggested that the conflict between pop-

FIGURE 18-3 Artificial selection in Brassica oleracea An enlarged terminal bud (the “head”) was selected in cabbage (lower left), flower clusters in broccoli (upper left) and cauliflower (middle right), axillary buds in brussels sprouts (bottom middle), leaves in collards (upper right) and kale (lower right), and stems in kohlrabi (middle).

ulation growth and food supply generates famine, disease, and war, which serve as inevitable brakes on population growth. Malthus’s idea that there is a strong and constant check on human population growth strongly influenced Darwin’s explanation of evolution. Darwin’s years of observing the habits of animals and plants had introduced him to the struggle for existence described by Malthus. It occurred to Darwin that in this struggle, inherited variations favorable to survival would tend to be preserved, whereas unfavorable ones would be eliminated. The result would be adaptation, an evolutionary modification that improves the chances of survival and reproductive success in a given environment. Eventually, the accumulation of modifications might result in a new species. Time was the only thing required for new species to originate, and the geologists of the era, including Lyell, had supplied evidence that Earth was indeed old enough to provide adequate time. Darwin had at last developed a workable scientific explanation of evolution, that of natural selection, in which better adapted organisms are more likely to survive and become the parents of the next generation. As a result of natural selection, the population changes over time; the frequency of favorable traits increases in successive generations, whereas less favorable traits become scarce or disappear. Darwin spent the next 20 years formulating his arguments for natural selection, accumulating an immense body of evidence to support his theory, and corresponding with other scientists. As Darwin was pondering his ideas, Alfred Russel Wallace (1823–1913), a British naturalist who studied the plants and animals of the Malay Archipelago for 8 years, was similarly struck by the diversity of species and the peculiarities of their distribution. He wrote a brief essay on this subject and sent it to Darwin, by then a world-renowned biologist, asking his opinion. Darwin rec-

3. Limits on population growth, or a struggle for existence. There is only so much food, water, light, growing space, and other resources available to a population, so organisms compete with one another for these limited resources. Because there are more individuals than the environment can support, not all survive to reproduce. Other limits on population growth include predators, disease organisms, and unfavorable weather conditions. 4. Differential reproductive success. Those individuals that have the most favorable combination of characteristics (those that make individuals better adapted to their environment) are more likely to survive and reproduce. Offspring tend to resemble their parents because the next generation inherits the parents’ genetically based traits. Successful reproduction is the key to natural selection: the best-adapted individuals produce the most offspring, whereas individuals that are less well adapted die prematurely or produce fewer or inferior offspring.

ognized his own theory and realized Wallace had independently arrived at the same conclusion—that evolution occurs by natural selection. Darwin’s colleagues persuaded him to present Wallace’s manuscript along with an abstract of his own work, which he had prepared and circulated to a few friends several years earlier. Both papers were presented in July 1858 at a London meeting of the Linnaean Society. Darwin’s monumental book, On the Origin of Species by Natural Selection; or, The Preservation of Favored Races in the Struggle for Life, was published in 1859. In 1870, Wallace’s book, Contributions to the Theory of Natural Selection, was published, 8 years after he returned from the Malay Archipelago.

Darwin proposed that evolution occurs by natural selection Darwin’s mechanism of evolution by natural selection consists of observations on four aspects of the natural world: variation; overproduction; limits on population growth, or a struggle for existence; and differential reproductive success.

© Biosphoto/Sauvenet Jany/Peter Arnold, Inc.

1. Variation. The individuals in a population exhibit variation (FIG. 18-4). Each individual has a unique combination of traits, such as size, color, ability to tolerate harsh environmental conditions, and resistance to certain parasites or infections. Some traits improve an individual’s chances of survival and reproductive success, whereas others do not. Remember that the variation necessary for evolution by natural selection must be inherited. Although Darwin recognized the importance to evolution of inherited variation, he did not know the mechanism of inheritance. 2. Overproduction. The reproductive ability of each species has the potential to cause its population to geometrically increase over time. A female frog lays about 10,000 eggs, and a female cod produces perhaps 40 million eggs! In each case, however, only about two offspring survive to reproduce. Thus, in every generation each species has the capacity to produce more offspring than can survive.

FIGURE 18-4 Genetic variation in emerald tree boas These snakes, all of the same species (Corallus caninus), were caught in a small section of forest in French Guiana. Many snake species exhibit considerable variation in their coloration and patterns.

Over time, enough changes may accumulate in geographically separated populations (often with slightly different environments) to produce new species. Darwin noted that the Galápagos finches appeared to have evolved in this way. The 14 species are closely related. All descended from a common ancestor—a single species that found its way from the South American mainland 2 million to 3 million years ago. (The closest genetic relatives of the Galápagos finches are small seed-eating birds known as grassquits that live in western South America.) During this 2-million to 3-million year period, the number of islands increased, the climate changed, and the plant life and food supply changed. The different islands of the Galápagos kept the finches isolated from one another, thereby allowing them to diverge into separate species in response to varying conditions (FIG. 18-5). Peter Grant, Rosemary Grant, and colleagues have documented natural selection in the Galápagos finches in their natural environment since the early 1970s. As an example of the evolutionary process in action, consider the sharp-beaked ground finch (Geospiza difficilis). This species lives on several different islands, and each population has evolved different beak shapes and sizes depending on the diet available on the island where it lives. We revisit the Galápagos finches in later chapters. Some longterm research by Peter Grant, Rosemary Grant, and colleagues on the microevolution of Galápagos finches when droughts affect the food supply is described in Chapter 19; these studies have demonstrated that environmental change can drive natural selection. Character displacement, an aspect of evolutionary ecology, is described in Galápagos finches in Chapter 54.

The modern synthesis combines Darwin’s theory with genetics One of the premises on which Darwin based his scientific theory of evolution by natural selection is that individuals transmit traits to the next generation. However, Darwin was unable to explain how this occurs or why individuals vary within a population. As discussed in Chapter 11, Gregor Mendel elucidated the basic patterns of inheritance. Darwin, who was a contemporary of Mendel, was apparently not acquainted with Mendel’s work. Indeed, the scientific community did not recognize Mendel’s work until the early part of the 20th century.

Miguel Castro/Photo Researchers, Inc.

© Images & Stories/Alamy

© Images & Stories/Alamy

© INTERFOTO/Alamy

(a) The cactus finch (Geospiza scandens), which feeds on the fleshy parts of cacti, such as their flowers, has a long, pointed beak.

(b) The large ground finch (Geospiza magnirostris) has an extremely heavy, nutcrackertype beak adapted for eating thick, hard-walled seeds.

(c) The warbler finch (Certhidia olivacea) has a slender beak for eating insects.

(d) The woodpecker finch (Camarhynchus pallidus) digs insects out of bark and crevices by using spines, twigs, or even dead leaves.

FIGURE 18-5 Animated Galápagos finches Darwin inferred that these birds are derived from a common ancestral population of seed-eating birds from South America. Variation in their beaks is the result of adaptation to the availability of different kinds of food.

Beginning in the 1930s and 1940s, biologists experienced a conceptual breakthrough when they combined the principles of Mendelian inheritance with Darwin’s theory of natural selection. The result was a unified explanation of evolution known as the modern synthesis. In this context, synthesis refers to combining parts of several theories to form a unified whole. Some of the founders of the modern synthesis were U.S. geneticist Theodosius Dobzhansky, British geneticist and statistician Ronald Fisher, British geneticist J. B. S. Haldane, British biologist Julian Huxley, U.S. biologist Ernst Mayr, U.S. paleontologist George Gaylord Simpson, U.S. botanist G. Ledyard Stebbins, and U.S. geneticist Sewell Wright. Today, the modern synthesis incorporates our expanding knowledge in genetics, which on its own makes an irrefutable case for evolution, systematics, paleontology, developmental biology, behavior, and ecology. The modern synthesis explains Darwin’s observation of variation among offspring in terms of mutation, or changes in DNA, such as nucleotide substitutions. Mutations provide the genetic variability on which natural selection acts during evolution. The modern synthesis, which emphasizes the genetics of populations as the central focus of evolution, has held up well since it was developed. It has dominated the thinking and research of biologists working in many areas and has resulted in an enormous accumulation of new discoveries that validate evolution by natural selection. Most biologists not only accept the basic principles of the modern synthesis but also try to better understand the causal processes of evolution. For example, what is the role of chance in evolution? How rapidly do new species evolve? These and other questions have arisen in part from a re-evaluation of the fossil record and in part from new discoveries in molecular aspects of inheritance. Such critical analyses are an integral part of the scientific process because they stimulate additional observation and experimentation, along with re-examination of previous evidence. Science is an ongoing process, and information obtained in the future may require modifications to certain parts of the modern synthesis.

We now consider one of the many evolutionary questions currently being addressed by biologists: the relative effects of chance and natural selection on evolution.

Biologists study the effect of chance on evolution Biologists have wondered whether we would get the same results if we were able to repeat evolution by starting with similar organisms exposed to similar environmental conditions. That is, would the same kinds of changes evolve, as a result of natural selection? Or would the organisms be quite different as a result of random events? Several recently reported examples of evolution in action suggest that chance may not be as important as natural selection, at least at the population level. A fruit fly species (Drosophila subobscura) native to Europe inhabits areas from Denmark to Spain. Biologists noted that the northern flies have larger wings than southern flies (FIG. 18-6). The same fly species was accidentally introduced to North America in the late 1970s. Ten years after its introduction, biologists determined that no statistically significant changes in wing size had occurred in the different regions of North America. However, 20 years after its introduction, the fruit flies in North America exhibited the same type of north–south wing changes as in Europe. (It is not known why larger wings evolve in northern areas and smaller wings in southern climates.) A study of the evolution of fishes known as sticklebacks in three coastal lakes of western Canada yielded intriguingly similar results to the fruit fly study. Molecular evidence indicates that when the lakes first formed several thousand years ago, they were populated with the same ancestral species. (Analysis of the mitochondrial DNA of sticklebacks in the three lakes supports the hypothesis of a common ancestor.) In each lake, the same two species have evolved from the common ancestral fish. One species is large and consumes invertebrates along the bottom of the lake, whereas the other species is smaller and consumes plankton at the lake’s surface.

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George Gilchrist

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FIGURE 18-6 Wing size in female fruit flies In Europe, female fruit flies (Drosophila subobscura) in northern countries have larger wings than flies in southern countries. Shown are two flies: one from Denmark (right) and the other from Spain (left). The same evolutionary pattern emerged in North America after the accidental introduction of D. subobscura to the Americas.

Members of the two species within a single lake do not interbreed with each other, but individuals of the larger species from one lake interbreed in captivity with individuals of the larger species from the other lakes. Similarly, smaller individuals from one lake interbreed in captivity with smaller individuals from the other lakes. In these examples, natural selection appears to be a more important agent of evolutionary change than chance. If chance were the most important factor influencing the direction of evolution, then fruit fly evolution would not have proceeded the same way on two different continents, and stickleback evolution would not have proceeded the same way in three different lakes. However, just because we have many examples of the importance of natural selection in evolution, it does not necessarily follow that random events should be discounted as a factor in directing evolutionary change. Proponents of the role of chance think that chance is more important in the evolution of major taxonomic groups (macroevolution) than in the evolution of populations (microevolution). It also may be that random events take place but their effects on evolution are harder to demonstrate than natural selection.

Review ■ ■ ■

What is natural selection? Why are only inherited variations important in the evolutionary process? What part of Darwin’s theory was he unable to explain? How does the modern synthesis fill this gap?

18.4 EVIDENCE FOR EVOLUTION ■ ■ LEARNING OBJECTIVES 5 Summarize the evidence for evolution obtained from the fossil record. 6 Define biogeography and describe how the distribution of organisms supports evolution.

Describe the evidence for evolution derived from comparative anatomy. Briefly explain how molecular biology and developmental biology provide insights into the evolutionary process. Give an example of how evolutionary hypotheses are tested experimentally.

A vast body of scientific evidence supports evolution, including observations from the fossil record, biogeography, comparative anatomy, molecular biology, and developmental biology. In addition, evolutionary hypotheses are increasingly being tested experimentally. Taken together, this evidence confirms the scientific theory that life unfolded on Earth by the process of evolution.

The fossil record provides strong evidence for evolution Perhaps the most direct evidence for evolution comes from the discovery, identification, and interpretation of fossils, which are the remains or traces typically left in sedimentary rock by previously existing organisms. (The term fossil comes from the Latin word fossilis, meaning “something dug up.”) Sedimentary rock forms by the accumulation and solidification of particles (pebbles, sand, silt, or clay) produced by the weathering of older rocks, such as volcanic rocks. The sediment particles, which are usually deposited on a riverbed, lake bottom, or the ocean floor, accumulate over time and exhibit distinct layers (FIG. 18-7). In an undisturbed rock sequence, the oldest layer is at the bottom, and upper layers are successively younger. The study of sedimentary rock layers, including their composition, arrangement, and similarity from one location to another, enables geologists to place events recorded in rocks in their correct sequence. The fossil record shows a progression from the earliest unicellular organisms to the many unicellular and multicellular organisms living today (see Table 21-1). The fossil record therefore demonstrates that life has evolved through time. To date, paleontologists (scientists who study extinct species) have described and named about 300,000 fossil species, and others are still being discovered. Although most fossils are preserved in sedimentary rock, some more recent remains have been exceptionally well preserved in bogs, tar, amber (ancient tree resin), or ice (FIG. 18-8). For example, the remains of a woolly mammoth deep-frozen in Siberian ice for more than 25,000 years were so well preserved that part of its DNA could be analyzed. Few organisms that die become fossils. The formation and preservation of a fossil require that an organism be buried under conditions that slow or prevent the decay process. This is most likely to occur if an organism’s remains are covered quickly by a sediment of fine soil particles suspended in water. In this way the remains of aquatic organisms may be trapped in bogs, mudflats, sandbars, or deltas. Remains of terrestrial organisms that lived on a floodplain may also be covered by waterborne sediments or, if the organism lived in an arid region, by windblown sand. Over time, the sediments harden to form sedimentary rock, and minerals usually replace the organism’s remains so that many details of its structure, even cellular details, are preserved. The fossil record is not a random sample of past life but instead is biased toward aquatic organisms and those living in the

© Alan Sirulnikoff/Photo Researchers, Inc.

evidence of the origin of new species from pre-existing species, including many transitional forms.

FIGURE 18-7 Exposed layers of sedimentary rock at the Burgess shale fossil bed This site, located in the Canadian Rockies, was formed about 500 million years ago when an avalanche of mud buried and preserved diverse and unusual marine animals.

few terrestrial habitats conducive to fossil formation. Relatively few fossils of tropical rainforest organisms have been found, for example, because their remains decay extremely rapidly on the forest floor, before fossils can develop. Another reason for bias in the fossil record is that organisms with hard body parts such as bones and shells are more likely to form fossils than are those with soft body parts. Also, rocks of different ages are unequally exposed at Earth’s surface; some rocks of certain ages are more accessible to paleontologists for fossil study than are rocks of other ages. Because of the nature of the scientific process, each fossil discovery represents a separate “test” of evolutionary theory. If any of the tests fail, the theory would have to be modified to fit the existing evidence. The verifiable discovery, for example, of fossil remains of modern humans (Homo sapiens) in Precambrian rocks, which are more than 570 million years old, would falsify evolutionary theory as currently proposed. However, Precambrian rocks examined to date contain only fossils of simple organisms, such as algae and small, soft-bodied animals, that evolved early in the history of life. The earliest fossils of H. sapiens with anatomically modern features do not appear in the fossil record until approximately 195,000 years ago (see Chapter 22). Fossils provide a record of ancient organisms and some understanding of where and when they lived. Using fossils of organisms from different geologic ages, scientists can sometimes infer the lines of descent (evolutionary relationships) that gave rise to modern-day organisms. In many instances, fossils provide direct

Transitional fossils document whale evolution Over the past century, biologists have found evidence suggesting that whales and other cetaceans (an order of marine mammals) evolved from land-dwelling mammals. During the 1980s and 1990s, paleontologists discovered several fossil intermediates in whale evolution that help document the whales’ transition from land to water. Fossils of Ambulocetus natans, a 50-million-year-old transitional form discovered in Pakistan, have many features of modern whales but also possess hind limbs and feet (FIG. 18-9a). (Modern whales do not have hind limbs, although vestigial pelvic and hind-limb bones persist. Vestigial structures are discussed later in the chapter.) The vertebrae of Ambulocetus’s lower back were very flexible, allowing the back to move dorsoventrally (up and down) during swimming and diving, as with modern whales. This ancient whale could swim but also moved about on land, perhaps as sea lions do today. Rodhocetus is a fossil whale found in more recent rocks in Pakistan (FIG. 18-9b). The vertebrae of Rodhocetus were even more flexible than those found in Ambulocetus. The flexible vertebrae allowed Rodhocetus a more powerful dorsoventral movement during swimming. Rodhocetus may have been totally aquatic. By 40 million years ago (mya), the whale transition from land to ocean was almost complete. Egyptian fossils of Basilosaurus show a whale with a streamlined body and front flippers for steering, like those of modern-day whales (FIG. 18-9c). Basilosaurus retained vestiges of its land-dwelling ancestors—a pair of reduced hind limbs that were disjointed from the backbone and probably not used in locomotion. Reduction in the hind limbs continued to the present. The modern blue whale has vestigial pelvis and femur bones embedded in its body (FIG. 18-9d). Various methods determine the age of fossils Because layers of sedimentary rock occur naturally in the sequence of their deposition, with the more recent layers on top of the older, earlier ones, most fossils are dated by their relative position in sedimentary rock. However, geologic events occurring after the rocks were initially formed have occasionally changed the relationships of some rock layers. Geologists identify specific sedimentary rocks not only by their positions in layers but also by features such as mineral content and by the fossilized remains of certain organisms, known as index fossils, that characterize a specific layer over large geographic areas. Index fossils are fossils of organisms that existed for a relatively short geologic time but were preserved as fossils in large numbers. With this information, geologists can arrange rock layers and the fossils they contain in chronological order and identify comparable layers in widely separated locations. Radioactive isotopes, also called radioisotopes, present in a rock provide a means to accurately measure its age (see Chapter 2). Radioisotopes emit invisible radiations. As radiation is emitted, the nucleus of a radioisotope changes into the nucleus of a different element; this process is known as radioactive decay. The radioactive nucleus of uranium-235, for example, decays into lead-207. Each radioisotope has its own characteristic rate of decay. The time required for one half of the atoms of a radioisotope to change into a different atom is known as its half-life (FIG. 18-10). Radioisotopes differ significantly in their half-lives. For example,

Bruce S. Lieberman/University of Kansas Kenneth Murray/Photo Researchers, Inc.

A. J. Copley/Visuals Unlimited

(a) Although some fossils contain traces of organic matter, all that remains in this fossil of a 500-millionyear-old jellyfish is an impression, or imprint, in the rock.

(d) A cast fossil of ancient echinoderms called crinoids formed when the crinoids decomposed, leaving a mold that later filled with dissolved minerals that hardened.

Scott Berner/Visuals Unlimited

Alfred Pasieka/Science Photo Library/Photo Researchers, Inc.

(b) Petrified wood from the Petrified Forest National Park in Arizona consists of trees that were buried and infiltrated with minerals. The logs were exposed by erosion of the mudstone layers in which they were buried.

(e) Dinosaur footprints, each 75 to 90 cm (2.5 to 3 ft) in length, provide clues about the posture, gait, and behavior of these extinct animals. (c) A 2-million-year-old insect fossil (a midge) was embedded in amber.

FIGURE 18-8 Fossils develop in different ways

100

(a) Ambulocetus natans, a transitional form between modern whale descendants and their terrestrial ancestors, possessed several recognizable whale features. It retained the hind limbs of its fourlegged ancestors, however.

Proportion of parent isotope remaining (%)

90 80 70

Decay of parent isotope

60 50 40 30 20 10 0 0

1

2 3 4 Time in half-lives

5

FIGURE 18-10 Animated Radioisotope decay

(b) The more recent Rodhocetus had flexible vertebrae that permitted a powerful dorsoventral movement during swimming.

(c) Basilosaurus was more streamlined and possessed tiny nonfunctional hind limbs.

(d) Balaenoptera, the modern blue whale, contains vestiges of pelvis and leg bones embedded in its body.

FIGURE 18-9 Fossil intermediates in whale evolution Biologists hypothesize that the ancestors of whales had four legs, as shown in these reconstructions of whale intermediates based on fossil evidence. Fossils indicate that ankle bones in whale intermediates match those found in present-day hippos, cows, pigs, and camels. Figures are not drawn to scale. (a–c: Adapted with permission from Futuyma, D. J., Science on Trial: The Case for Evolution, Fig. 2, pp. 260–261, Sinauer Associates, Sunderland, MA, 1995.)

the half-life of iodine-132 is only 2.4 hours, whereas the half-life of uranium-235 is 704 million years. The half-life of a particular radioisotope is constant and does not vary with temperature, pressure, or any other environmental factor. The age of a fossil in sedimentary rock is usually estimated by measuring the relative proportions of the original radioisotope and its decay product in volcanic rock intrusions that penetrate the sediments. For example, the half-life of potassium-40 is 1.3 billion years, meaning that in 1.3 billion years half of the radioactive potassium will have decayed into its decay product, argon-40. The

At time zero, the sample is composed entirely of the radioisotope and the radioactive clock begins ticking. After one half-life, only 50% of the original radioisotope remains. During each succeeding half-life, half of the remaining radioisotope is converted to decay product(s).

radioactive clock begins ticking when the magma solidifies into volcanic rock. The rock initially contains some potassium but no argon. Because argon is a gas, it escapes from hot rock as soon as it forms, but when potassium decays in rock that has cooled and solidified, the argon accumulates in the crystalline structure of the rock. If the ratio of potassium-40 to argon-40 in the rock being tested is 1:1, the rock is 1.3 billion years old. Several radioisotopes are commonly used to date fossils. Three of these are potassium-40 (half-life 1.3 billion years), uranium-235 (half-life 704 million years), and carbon-14 (half-life 5730 years). Radioisotopes other than carbon-14 are used to date the rock in which fossils are found, whereas carbon-14 is used to date the carbon remains of anything that was once living, such as wood, bones, and shells. Whenever possible, the age of a fossil is independently verified using two or more different radioisotopes. Carbon-14, which is continuously produced in the atmosphere from nitrogen-14 (by cosmic radiation), subsequently decays to nitrogen-14. Because the formation and the decay of carbon-14 occur at constant rates, the ratio of carbon-14 to carbon-12 (a more abundant, stable isotope of carbon) is constant in the atmosphere. Organisms absorb carbon from the atmosphere either directly (by photosynthesis) or indirectly (by consuming photosynthetic organisms). Because each organism absorbs carbon from the atmosphere, its ratio of carbon-14 to carbon-12 is the same as that in the atmosphere. When an organism dies, however, it no longer absorbs carbon, and the proportion of carbon-14 in its remains declines as carbon-14 decays to nitrogen-14. Because of its relatively short halflife, carbon-14 is useful for dating fossils that are 50,000 years old or less. It is particularly useful for dating archaeological sites.

The distribution of plants and animals supports evolution The study of the past and present geographic distribution of organisms is called biogeography. The geographic distribution of

organisms affects their evolution. Darwin was interested in biogeography, and he considered why the species found on ocean islands tend to resemble species of the nearest mainland, even if the environment is different. He also observed that species on ocean islands do not tend to resemble species on islands with similar environments in other parts of the world. Darwin studied the plants and animals of two sets of arid islands—the Cape Verde Islands, nearly 640 km (400 mi) off western Africa, and the Galápagos Islands, about 960 km (600 mi) west of Ecuador, South America. On each group of islands, the plants and terrestrial animals were indigenous (native), but those of the Cape Verde Islands resembled African species and those of the Galápagos resembled South American species. The similarities of Galápagos species to South American species were particularly striking considering that the Galápagos Islands are dry and rocky and the nearest part of South America is humid and has a lush tropical rain forest. Darwin concluded that species from the neighboring continent migrated or were carried to the islands, where they subsequently adapted to the new environment and, in the process, evolved into new species. If evolution were not a factor in the distribution of species, we would expect to find a given species everywhere that it could survive. However, the actual geographic distribution of organisms makes sense in the context of evolution. For example, Aus-

KEY POINT

tralia, which has been a separate landmass for millions of years, has distinctive organisms. Australia has populations of egg-laying mammals (monotremes) and pouched mammals (marsupials) not found anywhere else. Two hundred million years ago, Australia and the other continents were joined in a major landmass. Over the course of millions of years, the Australian continent gradually separated from the others. The monotremes and marsupials in Australia continued to thrive and diversify. The isolation of Australia also prevented placental mammals, which arose elsewhere at a later time, from competing with its monotremes and marsupials. In other areas of the world where placental mammals evolved, most monotremes and marsupials became extinct. We now consider how Earth’s dynamic geology has affected biogeography and evolution. Biogeography and evolution are related to Earth’s geologic history In 1915, the German scientist Alfred Wegener, who had noted a correspondence between the geographic shapes of South America and Africa, proposed that all the landmasses had at one time been joined into one huge supercontinent, which he called Pangaea (FIG. 18-11a). He further suggested that Pangaea had subsequently broken apart and that the various landmasses had separated in a process known as continental drift. Wegener did not know of any

Geologists hypothesize that the breakup of Pangaea is only the latest in a series of continental breakups and collisions that have taken place since early in Earth’s history.

N. America

Laurasia Pangaea

S. America

Gondwana

Asia Europe Africa India

Australia Antarctica

(a) The supercontinent Pangaea, about 240 mya (Triassic period).

(b) Breakup of Pangaea into Laurasia (Northern Hemisphere) and Gondwana (Southern Hemisphere) began about 180 mya. The landmasses looked this way about 120 mya (Cretaceous period).

(c) Further separation of landmasses, 60 mya (early Paleogene period). Note that Europe and North America were still joined and that India was a separate landmass.

N. America

Eurasia Africa

S. America Australia

Antarctica

(d) The continents today.

FIGURE 18-11 Animated Continental drift

mechanism that could have caused continental drift, so his idea, although debated initially, was largely ignored. In the 1960s, scientific evidence provided the explanation for continental drift. Earth’s crust is composed of seven large plates (plus a few smaller ones) that float on the mantle, which is the mostly solid layer of Earth lying beneath the crust and above the core. (Because of its hotter temperature, the solid rock of the mantle is more plastic than the solid rock of the crust above it.) The landmasses are situated on some of these plates. As the plates move, the continents change their relative positions (FIGS. 18-11b, c, and d). The movement of the crustal plates is termed plate tectonics. Any area where two plates meet is a site of intense geologic activity. Earthquakes and volcanoes are common in such a region. Both San Francisco, noted for its earthquakes, and the Mount Saint Helens volcano are situated where two plates meet. If landmasses lie on the edges of two adjacent plates, mountains may form. The Himalayas formed when the plate carrying India rammed into the plate carrying Asia. When two plates grind together, one of them is sometimes buried under the other in a process known as subduction. When two plates move apart, a ridge of lava forms between them. The Atlantic Ocean is increasing in size because of the expanding zone of lava along the Mid-Atlantic Ridge, where two plates are separating. Knowledge that the continents were at one time connected and have since drifted apart is useful in explaining certain aspects of biogeography (FIG. 18-12). Likewise, continental drift has played a major role in the evolution of different organisms. When Pangaea originally formed during the late Permian period, it brought together terrestrial species that had evolved separately from one another, leading to competition and some extinction. Marine life was adversely affected, in part because, with the continents joined as one large mass, less coastal area existed. (Shallow coastal areas contain a greater diversity of marine species than deep-water environments.) Pangaea separated into several landmasses approximately 180 mya. As the continents began to drift apart, populations became geographically isolated, were exposed to different environmental conditions, and began to diverge along separate evolutionary pathways. As a result, the plants, animals, and other organisms of previously connected continents—South America and Africa, for example—differ. Continental drift also caused gradual changes in ocean and atmospheric currents that have profoundly influenced the biogeography and evolution of organisms. (Biogeography is discussed further in Chapter 56.)

Comparative anatomy of related species demonstrates similarities in their structures Comparing the structural details of features found in different but related organisms reveals a basic similarity. Such features that are derived from the same structure in a common ancestor are termed homologous features; the condition is known as homology. For example, consider the limb bones of mammals. A human arm, a cat forelimb, a whale front flipper, and a bat wing, although quite different in appearance, have strikingly similar arrangements of bones, muscles, and nerves. FIGURE 18-13 shows a comparison of their skeletal structures. Each has a single bone (the humerus) in

KEY POINT

Knowledge that the continents were united at one time explains the unique distribution of certain fossil plants and animals.

(a) Cynognathus was a carnivorous reptile found in Triassic rocks in South America and Africa.

(b) Lystrosaurus was a herbivorous reptile found in Triassic rocks in Africa, India, and Antarctica.

Cynognathus

Lystrosaurus

Africa India South America Australia Antarctica

Mesosaurus

(c) Mesosaurus was a freshwater reptile found in Permian rocks in South America and Africa.

Glossopteris

(d) Glossopteris was a seed-bearing tree found in Permian rocks in South America, Africa, India, Antarctica, and Australia.

Adapted from E. H. Colbert, Wandering Lands and Animals, Hutchinson, London, 1973.

FIGURE 18-12 Distribution of fossils on continents that were joined during the Permian and Triassic periods (286 mya to 213 mya)

the part of the limb nearest the trunk of the body, followed by the two bones (radius and ulna) of the forearm, a group of bones (carpals) in the wrist, and a variable number of digits (metacarpals and phalanges). This similarity is particularly striking because arms, forelimbs, flippers, and wings are used for different types of locomotion, and there is no overriding mechanical reason for them to be so similar structurally. Similar arrangements of parts of the forelimb are evident in ancestral birds, reptiles, and amphibians and even in the first fishes that came out of water onto land hundreds of millions of years ago (see Fig. 32-17).

HUMAN CAT

WHALE

BAT

Humerus Radius

Ulna

Humerus

Carpal Radius

5

Metacarpal 4

Ulna Carpal

1

Radius Ulna

1 5

Carpal Metacarpal

1

1 Phalanges 5 4

54

3 3

2

Phalanges

2

4

2 3

3 2 Tendril

FIGURE 18-13 Animated Structural homology in vertebrates

Spine

The human arm, cat forelimb, whale flipper, and bat wing have a basic underlying similarity of structure because they are derived from a common ancestor. The five digits are numbered in each drawing.

Leaflet

Leaves provide an example of homology in plants. In many plant species, leaves have been modified for functions other than photosynthesis. A cactus spine and a pea tendril, although quite different in appearance, are homologous because both are modified leaves (FIG. 18-14). The spine protects the succulent stem tissue of the cactus, whereas the tendril, which winds around a small object once it makes contact, helps support the climbing stem of the pea plant. Such modifications in organs used in different ways are the expected outcome of a common evolutionary origin. The basic structure present in a common ancestor was modified in different ways for different functions as various descendants subsequently evolved. Not all species with “similar” features have descended from a recent common ancestor, however. Sometimes similar environmental conditions result in the evolution of similar adaptations. Such independent evolution of similar structures in distantly related organisms is known as convergent evolution. Aardvarks, anteaters, and pangolins are an excellent example of convergent evolution (FIG. 18-15). They resemble one another in lifestyle and certain structural features. All have strong, sharp claws to dig open ant and termite mounds and elongated snouts with long, sticky tongues to catch these insects. Yet aardvarks, anteaters, and pangolins evolved from three distantly related orders of mammals. (See Figure 32-28, which shows several examples of convergent evolution in placental and marsupial mammals.)

Leaf petiole

(a)

Stipule Stem

(b)

FIGURE 18-14 Homology in plants (a) The spines of the fishhook cactus (Ferocactus wislizenii) are modified leaves, as are (b) the tendrils of the garden pea (Pisum sativum). Leaves of the garden pea are compound, that is, they are divided into smaller components called leaflets. The terminal leaflets are modified into tendrils that are frequently branched.

Structurally similar features that are not homologous but have similar functions that evolved independently in distantly related organisms are said to be homoplastic features. Such similarities in different species that are independently acquired by convergent evolution and not by common descent are called homoplasy.2

2

An older, less precise term that some biologists still use for nonhomologous features with similar functions is analogy.

Kjell B. Sandved/Visuals Unlimited Gunter Ziesler/Peter Arnold, Inc.

(a) The aardvark (Orycteropus afer) is native to central, southern, and eastern Africa.

(b) A giant anteater (Myrmecophaga tridactyla) at a termite mound. The anteater is native to Latin America, from southern Mexico to northern Argentina.

Spines, which are modified leaves, and thorns, which are modified stems, are an example of homoplasy in plants. Spines and thorns resemble one another superficially but are homoplastic features that evolved independently to solve the common need for protection from herbivores (FIG. 18-16). Like homology, homoplasy offers crucial evidence of evolution. Homoplastic features are of evolutionary interest because they demonstrate that organisms with separate ancestries may adapt in similar ways to similar environmental demands. Comparative anatomy also reveals the existence of vestigial structures. Many organisms contain organs or parts of organs that are seemingly nonfunctional and degenerate, often undersized or lacking some essential part. Vestigial structures are remnants of more developed structures that were present and functional in ancestral organisms. In the human body, more than 100 structures are considered vestigial, including the coccyx (fused tailbones), third molars (wisdom teeth), and the muscles that move our ears. Whales and pythons have vestigial hind-limb bones (FIG. 18-17); pigs have vestigial toes that do not touch the ground; wingless birds such as the kiwi have vestigial wing bones; and many blind, burrowing, or cave-dwelling animals have nonfunctioning, vestigial eyes. The occasional presence of a vestigial structure is to be expected as a species adapts to a changing mode of life. Some structures become much less important for survival and may end up as vestiges. When a structure no longer confers a selective advantage, it may become smaller and lose much or all of its function with the passage of time. Because the presence of the vestigial structure is usually not harmful to the organism, however, selective pressure for completely eliminating it is weak, and the vestigial structure is found in many subsequent generations.

Mandal Ranjit/Photo Researchers, Inc.

Molecular comparisons among organisms provide evidence for evolution

(c) The pangolin (Manis crassicaudata) is native to Africa and to southern and southeastern Asia.

FIGURE 18-15 Convergent evolution Three distantly related mammals adapted independently to eat ants and termites in similar grassland/forest environments in different parts of the world.

For example, the wings of various distantly related flying animals, such as insects and birds, resemble one another superficially; they are homoplastic features that evolved over time to meet the common function of flight, although they differ in more fundamental aspects. Bird wings are modified forelimbs supported by bones, whereas insect wings may have evolved from gill-like appendages present in the aquatic ancestors of insects.

Fossils, biogeography, and comparative anatomy provided Darwin with important clues about the evolutionary history of life. Today, similarities and differences in the biochemistry and molecular biology of various organisms provide additional compelling evidence for evolutionary relationships. Molecular evidence for evolution includes the universal genetic code and the conserved sequences of amino acids in proteins and of nucleotides in DNA. The genetic code is virtually universal Organisms owe their characteristics to the types of proteins they possess, which in turn are determined by the sequence of nucleotides in their messenger ribonucleic acid (mRNA), as specified by the order of nucleotides in their DNA. Evidence that all life is related comes from the fact that all organisms use a genetic code that is virtually identical. (There is some minor variation in the genetic code. For example, some mitochondria have several deviations from the standard code.) Recall from Chapter 13 that the genetic code specifies a triplet (a sequence of three nucleotides in DNA) that codes for a particular codon (a sequence of three nucleotides in mRNA). The codon then codes for a particular amino acid in a polypeptide chain. For example, “AAA” in DNA codes for “UUU” in mRNA, which codes for the amino acid phenylalanine in organisms as diverse as

J. D. Cunningham/Visuals Unlimited

E. R. Degginger/Animals Animals

or so. Moreover, the recent explosion in DNA sequencShoot ing information (more than (develops 1000 species have now had from axillary their genomes sequenced) bud) is beginning to provide a wealth of evolutionary data as these genomes are compared. Spine Thorn (develops Sequence-based relationships (midrib from axillary bud) Leaf scar generally agree with earlier of leaf) studies, which based evolutionary relationships on similarities in structure among living organisms and on fossil data of extinct organisms. Investigations of the sequence of amino acids in pro(a) A spine of Japanese barberry (Berberis (b) Thorns of downy hawthorn (Crataegus mollis) thunbergii) is a modified leaf. (In this example, are modified stems that develop from axillary buds. teins that play the same roles the spine is actually the midrib of the original in many species have revealed leaf, which has been shed.) both great similarities and certain specific differences. Even organisms that are only remotely FIGURE 18-16 Homoplasy in plants related share some proteins, such as cytochrome c, which is part of the electron transport chain in aerobic respiration. To survive, all aerobic organisms need a respiratory protein with the same basic shrimp, humans, bacteria, and tulips. In fact, “AAA” codes for phestructure and function as the cytochrome c of their common annylalanine in all organisms examined to date. cestor. Consequently, not all amino acids that confer the structural The universality of the genetic code—no other code has been and functional features of cytochrome c are free to change. Any found in any organism—is convincing evidence that all organisms mutations that changed the amino acid sequence at structurally arose from a common ancestor. The genetic code has been mainimportant sites of the cytochrome c molecule would have been tained and transmitted through all branches of the evolutionary tree harmful, and natural selection would have prevented such mutasince its origin in some extremely early (and successful) organism. tions from being passed to future generations. However, in the course of the long, independent evolution of different organisms, Proteins and DNA contain a record of evolutionary change mutations have resulted in the substitution of many amino acids at Researchers have carried out thousands of comparisons of protein less important locations in the cytochrome c molecule. The greater and DNA sequences from various species during the past 30 years the differences in the amino acid sequences of their cytochrome c molecules, the longer it has been since two species diverged. Because a protein’s amino acid sequences are coded in DNA, the differences in amino acid sequences indirectly reflect the nature and number of underlying DNA base-pair changes that must have occurred during evolution. Of course, not all DNA codes for proteins (witness introns and transfer RNA genes). DNA sequencing—that is, determining the order of nucleotide bases in DNA—of both protein-coding DNA and nonprotein-coding DNA—is useful in determining evolutionary relationships. Generally, the more closely species are considered related on the basis of other scientific evidence, the greater the percentage of nucleotide sequences that their DNA molecules have in common. By using the DNA sequence data in FIGURE 18-18, for example, you can conclude that the closest living relative of humans is the chimpanzee (because its DNA has the lowest percentage of differences in the sequence examined). (Primate evolution is discussed in Chapter 22.) In some cases, molecular evidence challenges traditional evo(a) An African rock python (b) Close-up of part of a python lutionary ideas that were based on structural comparisons among (Python sebae). skeleton showing the hind-limb bones. living species and/or on studies of fossil skeletons. Consider eventoed hoofed mammals such as pigs, camels, deer, antelope, cattle, and hippos. Whales lack toes and were traditionally not classified FIGURE 18-17 Vestigial structures as close relatives of hoofed mammals. All pythons have remnants of hind-limb bones embedded in their bodies.

Primate species compared to humans

changes—how genes are switched on and off during development—help explain the diversity of form in species with similar Chimpanzee genes. Mutations in genes that regulate development often result in dramatically different structures (see Fig. 17-12). Gorilla Increasingly, developmental biology, particularly at the molecular level, is proOrangutan viding answers to such questions as how snakes became elongated and lost their limbs. In many cases, evolutionary changes Gibbon such as the loss of limbs in snakes occur as a result of changes in genes that regulate the Rhesus monkey orderly sequence of events that occurs during development. In pythons, for example, the loss of forelimbs and elongation of the Spider monkey body are linked to mutations in several Hox genes that affect the expression of body patTarsier terns and limb formation in a wide variety of animals. (See the discussion of Hox gene 5 10 15 20 25 clusters in Chapter 17.) The hind limbs may Percent divergence in noncoding not develop because python embryonic sequence of β-globin gene tissue does not respond to internal signals that trigger leg elongation. Developmental geneticists at Harvard Medical School and Princeton University are FIGURE 18-18 Differences in DNA nucleotide sequences as evidence of evolutionary studying the developmental basis for the difrelationships ferent beak shapes of the Galápagos finches. Comparing the same gene in different organisms provides a window into evolution. Here the differThey have determined that a gene that codes ences in the noncoding region of the b-globin gene are compared between humans and other primates. Evolutionary biologists are beginning to expand such studies from comparing one or several genes to for an important signaling molecule, bone comparing entire genomes. morphogenic protein 4 (BMP4), affects the development of the birds’ craniofacial skeletons. The gene for BMP4 is turned on earlier FIGURE 18-19 depicts a cladogram, or phylogenetic tree, based in development and has a higher level of expression in finch species on molecular data for whales and selected hoofed mammals. Such with larger, thicker beaks than in finches with smaller beaks. cladograms—diagrams showing lines of descent—can be derived Scientific evidence overwhelmingly demonstrates that defrom differences in a given DNA nucleotide sequence. This diagram velopment in different animals is controlled by the same kinds of indicates that whales are more closely related to hippos than to any genes; these genetic similarities in a wide variety of organisms reother hoofed mammal. The branches representing whales and hipflect a shared evolutionary history. For example, vertebrates have pos probably diverged relatively recently because of the close simisimilar patterns of embryological development that indicate they larity of DNA sequences in these species. In contrast, camels, which share a common ancestor. All vertebrate embryos have segmented have DNA sequences that are less similar to those of whales, dimuscles, pharyngeal (throat) pouches, a tubular heart without verged much earlier. The molecular evidence indicates that whales left and right sides, a system of arteries known as aortic arches in and hippos share a recent common ancestor. the pharyngeal region, and many other shared features. All these The molecular evidence linking whales and even-toed hoofed structures are necessary and functional in the developing fish. The mammals has led to a reexamination of early fossil whales. Like small, segmented muscles of the fish embryo give rise to the segtoday’s hoofed mammals, early fossil whales had an even number mented muscles used by the adult fish in swimming. The pharynof toes on their appendages. Fossil whales also had the same type geal pouches break through to the surface as gill slits. The adult fish of specialized ankle bone (called a double pulley) as hippos and heart remains undivided and pumps blood forward to the gills that other hoofed mammals. develop in association with the aortic arches. Because none of these embryonic features persists in the adults of reptiles, birds, or mammals, why are these fishlike strucDevelopmental biology helps tures present in their embryos? Evolution is a conservative process, unravel evolutionary patterns and natural selection builds on what has come before rather than As early as 1975, biologists were suggesting that regulatory changes starting from scratch. The evolution of new features often does not in gene expression, particularly of genes involved in development, require the evolution of new developmental genes but instead dewere responsible for many differences between closely related pends on a modification in developmental genes that already exist species. Today much evidence supports the idea that regulatory (see discussion of pre-adaptations in Chapter 20). Terrestrial ver-

Chicken

Human

1 day

3 days

32 days

3 days

12 days

56 days

Common ancestor of hippos and whales

Common ancestor of even-toed hoofed mammals and cetaceans Adapted from Nikaido, M., et al., “Phylogenetic Relationships among Cetartiodactyls Based on Insertions of Short and Long Interspersed Elements: Hippopotamuses Are the Closest Extant Relatives of Whales,” Proceedings of the National Academy of Sciences, Vol. 96, Aug. 31, 1999.

FIGURE 18-19 Cladogram of whales and their closest living relatives DNA sequence differences among selected mammals suggest that hoofed mammals such as hippos and giraffes are the closest living relatives of whales. The hippopotamus is probably the closest living relative of whales and other cetaceans. The nodes (circles) represent branch points where a species splits into two or more lineages. (Ruminants are mammals such as cows, sheep, and giraffes that have a multichambered stomach and chew regurgitated plant material to make it more digestible.)

tebrates are thought to have evolved from fishlike ancestors; therefore, they share some of the early stages of development still found in fishes today. The accumulation of genetic changes over time in these vertebrates has modified the basic body plan laid out in fish development (FIG. 18-20).

Evolutionary hypotheses are tested experimentally Increasingly, biologists are designing imaginative experiments, often in natural settings, to test evolutionary hypotheses. David Reznick from the University of California at Santa Barbara and John Endler from James Cook University in Australia have studied evolution in guppy populations in Venezuela and in Trinidad, a small island in the southern Caribbean.

FIGURE 18-20 Development of fish fins, chicken wings, and human limbs Fish, chickens, and humans are vertebrates with strikingly similar genes. Although the early embryos of these organisms are much alike, the areas colored orange follow different developmental pathways, resulting in fins, wings, or limbs. (Figures are not to scale.)

Reznick and Endler observed that different streams have different kinds and numbers of fishes that prey on guppies. Predatory fishes that prey on larger guppies are present in all streams at lower elevations; these areas of intense predation pressure are known as high-predation habitats. Predators are often excluded from tributaries or upstream areas by rapids and waterfalls. The areas above such barriers are known as low-predation habitats because they contain only one species of small predatory fish that occasionally eats smaller guppies. Differences in predation are correlated with many differences in the guppies, such as male coloration, behavior, and attributes known as life history traits (discussed in more detail in Chapter 53). These traits include age and size at sexual maturity, the number of offspring per reproductive event, the size of the offspring, and the frequency of reproduction. For example, guppy adults are larger in streams found at higher elevations and smaller in streams found at lower elevations. Do predators actually cause these differences to evolve? Reznick and colleagues tested this evolutionary hypothesis by conducting field experiments in Trinidad. Taking advantage of waterfalls that prevent the upstream movement of guppies, guppy predators, or both, they moved either guppies or guppy predators over such barriers. For example, guppies from a high-predation habitat were introduced into a low-predation habitat by moving them over a barrier waterfall into a section of stream that was free of guppies and large predators. The only fish species that lived in this section of stream before the introduction was the small predatory fish.

John Burgoyne/National Geographic Image Collection

Fish

Cetacean

Hippopotamus

This branching diagram, called a cladogram, shows hypothetical evolutionary relationships. The organisms depicted here share a common ancestor.

Ruminant

Pig

Camel

KEY POINT

KEY EXPERIMENT QUESTION: Can natural selection be observed in a natural population?

HYPOTHESIS: A natural population will respond adaptively to environmental change. EXPERIMENT: Male and female guppies from a stream in which the predators preferred large adult guppies as prey (brown bars) were transferred to a stream in which the predators preferred juveniles and small adults. 200 Average weight of adult guppy (mg)

Eleven years later, the researchers captured adult females from the introduction site (low-predation habitat) and the control site below the barrier waterfall (high-predation habitat). They bred these females in their laboratory and compared the life history traits of succeeding generations. The descendants of guppies introduced into the low-predation habitat matured at a larger size than did the descendants of guppies from the control site below the waterfall (FIG. 18-21). They also produced fewer, but larger, offspring. The introduced fish populations had evolved to have life histories similar to those of fishes typically found in such low-predation habitats. Similar studies have demonstrated that predators have played an active role in the evolution of other traits, such as the average number of offspring produced during the lifetime of an individual female (fecundity), male coloration, and behavior. Rapid evolution on a scale of years has also been observed in organisms as diverse as marine snails, mussels, soapberry bugs, mayflies, anole lizards, ’i’iwis (scarlet honeycreepers), and wild rabbits. These and other experiments and observations demonstrate not only that evolution is real but also that it is occurring now, driven by selective environmental forces, such as predation, that can be experimentally manipulated. Darwin incorrectly assumed evolution to be so gradual that humans cannot observe it. As Jonathan Weiner, author of The Beak of the Finch: A Story of Evolution in Our Time, notes, “Darwin did not know the strength of his own theory. He vastly underestimated the power of natural selection. Its action is neither rare nor slow. It leads to evolution daily and hourly, all around us, and we can watch.”

■

■

■

■ ■

Average initial weight of guppies (control)

80

40

Males

Females

Average weight of guppies 18 generations after being transferred to stream with different selective predation (experimental)

RESULTS AND CONCLUSION: After 11 years, the descendants of the transferred guppies (pink bars) were measurably larger than their ancestors, indicating that larger guppies had a selective advantage in the new environment.

How do scientists date fossils? How can we explain that fossils of Mesosaurus, an extinct reptile that could not swim across open water, are found in the southern parts of both Africa and South America? How do homologous and homoplastic features provide evidence of evolution? How does developmental biology provide evidence of a common ancestry for vertebrates as diverse as reptiles, birds, pigs, and humans? How do predator preferences drive the evolution of size in guppies?

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120

0

Review ■

160

Data used with permission from Reznick, D. N., et al., “Evaluation of the Rate of Evolution in Natural Populations of Guppies [Poecilia reticulata],” Science, Vol. 275, Mar. 28, 1997.

FIGURE 18-21 D. N. Reznick’s experiment in guppy

evolution

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

18.1 (page 392)

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1 Define evolution. ■

Evolution is the accumulation of inherited changes within a population over time. Evolution is the unifying concept of biology because it links all fields of the life sciences into a coherent body of knowledge.

18.2 (page 392) 2 Discuss the historical development of evolutionary theory. ■

Jean Baptiste de Lamarck was the first scientist to propose that organisms undergo change over time as a result of some natural phenomenon rather than divine intervention. Lamarck thought organisms were endowed with a vital force that drove them to change toward greater complexity over time. He thought organisms could pass traits acquired during their lifetimes to their offspring.

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Charles Darwin’s observations while voyaging on the HMS Beagle were the basis for his evolutionary theory. Darwin tried to explain the similarities between animals and plants of the arid Galápagos Islands and the humid South American mainland. Darwin was influenced by artificial selection, in which breeders develop many varieties of domesticated plants and animals in just a few generations. Darwin applied Thomas Malthus’s ideas on the natural increase in human populations to natural populations. Darwin was influenced by the idea that Earth was extremely old, an idea promoted by Charles Lyell and other geologists.

18.3 (page 393) 3 Explain the four premises of evolution by natural selection as proposed by Charles Darwin. Charles Darwin and Alfred Wallace independently proposed evolu■ tion by natural selection, which is based on four observations.

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First, genetic variation exists among the individuals in a population. Second, the reproductive ability of each species causes its populations to have the potential to geometrically increase in number over time. Third, organisms compete with one another for the resources needed for life, such as food, living space, water, and light. Fourth, offspring with the most favorable combination of inherited characteristics are most likely to survive and reproduce, passing those genetic characteristics to the next generation. Natural selection results in adaptations, evolutionary modifications that improve the chances of survival and reproductive success in a particular environment. Over time, enough changes may accumulate in geographically separated populations to produce new species.

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mologous features derive from the same structure in a common ancestor. Evolutionary affinities exist among the organisms that have homologous features. Homoplastic features evolved independently to have similar functions in distantly related organisms. Homoplastic features demonstrate convergent evolution, in which organisms with separate ancestries adapt in similar ways to comparable environmental demands.

Learn more about the Galápagos Islands and the Galápagos Island finches by clicking on the figures in CengageNOW.

4 Compare the modern synthesis with Darwin’s original evolutionary theory. The modern synthesis combines Darwin’s evolutionary theory by natural selection with modern genetics to explain why individuals in a population vary and how species adapt to their environment. ■ Mutation provides the genetic variability that natural selection acts on during evolution. ■

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18.4 (page 397) 5 Summarize the evidence for evolution obtained from the fossil record. ■

Direct evidence of evolution comes from fossils, the remains or traces of ancient organisms. Layers of sedimentary rock normally occur in their sequence of deposition, with the more recent layers on top of the older, earlier ones. Index fossils characterize a specific layer over large geographic areas. Radioisotopes present in a rock provide a way to accurately measure the rock’s age. Learn more about fossil intermediates by clicking on the figure in CengageNOW.

6 Define biogeography and describe how the distribution of organisms supports evolution. Biogeography, the geographic distribution of organisms, affects their evolution. Areas that have been separated from the rest of the world for a long time contain organisms that have evolved in isolation and are therefore unique to those areas. ■ At one time the continents were joined to form a supercontinent. Continental drift, which caused the various landmasses to break apart and separate, has played a major role in evolution. ■

Watch continental drift by clicking on the figure in CengageNOW.

7 Describe the evidence for evolution derived from comparative anatomy. ■

Homologous features have basic structural similarities, even though the structures may be used in different ways because ho-

Vestigial structures are nonfunctional or degenerate remnants of structures that were present and functional in ancestral organisms. Structures occasionally become vestigial as species adapt to different modes of life. Learn more about vestigial structures by clicking on the figure in CengageNOW.

8 Briefly explain how molecular biology and developmental biology provide insights into the evolutionary process. ■ Molecular evidence for evolution includes the universal genetic code and the conserved sequences of amino acids in proteins and of nucleotides in DNA. ■ Evolutionary changes are often the result of mutations in genes that affect the orderly sequence of events during development. Development in different animals is controlled by the same kinds of genes, which indicates these animals have a shared evolutionary history. ■ The accumulation of genetic changes since organisms diverged, or took separate evolutionary pathways, has modified the pattern of development in more complex vertebrate embryos. 9 Give an example of how evolutionary hypotheses are tested experimentally. ■ Reznick and Endler have studied the effects of predation intensity on the evolution of guppy populations in the laboratory and in nature. Such experiments are a powerful way for investigators to test the underlying processes of natural selection.

T E S T YO U R U N D E R S TA N D I N G 1. Evolution is based on which of the following concepts? (a) organisms share a common origin (b) over time, organisms have diverged from a common ancestor (c) an animal’s body parts can change over its lifetime, and these acquired changes are passed to the next generation (d) a and b (e) a, b, and c

3. Charles Darwin proposed that evolution could be explained by the differential reproductive success of organisms that resulted from their naturally occurring variation. Darwin called this process (a) coevolution (b) convergent evolution (c) natural selection (d) artificial selection (e) homoplasy

2. Evolution is the accumulation of genetic changes within _____ over time. (a) individuals (b) populations (c) communities (d) a and b (e) a and c

4. Which of the following is not part of Darwin’s mechanism of evolution? (a) differential reproductive success (b) variation

in a population (c) inheritance of acquired (nongenetic) traits (d) overproduction of offspring (e) struggle for existence 5. The modern synthesis (a) is based on the sequence of fossils in rock layers (b) uses genetics to explain the source of hereditary variation that is essential to natural selection (c) was first proposed by ancient Greek scholars (d) considers the influence of the geographic distribution of organisms on their evolution (e) is reinforced by homologies that are explained by common descent 6. The evolution of beak size in the various species of Galápagos finches is associated with their (a) songs (b) diets (c) body sizes (d) predators (e) none of the preceding 7. The fossil record (a) usually occurs in sedimentary rock (b) sometimes appears fragmentary (c) is relatively complete for tropical rainforest organisms but incomplete for aquatic organisms (d) a and b (e) a, b, and c 8. In _____, the selecting agent is the environment, whereas in _____, the selecting agent is humans. (a) natural selection; con-

vergent evolution (b) mutation; artificial selection (c) homoplasy; homology (d) artificial selection; natural selection (e) natural selection; artificial selection 9. Aardvarks, anteaters, and pangolins are only distantly related but are similar in structure and form as a result of (a) homology (b) convergent evolution (c) biogeography (d) vestigial structures (e) artificial selection 10. The species of the Galápagos Islands (a) are similar to those on other islands at the same latitude (b) are similar to those on the South American mainland (c) are identical to those on other islands at the same latitude (d) are identical to those on the South American mainland (e) are similar to those on both the African and South American mainlands 11. Changes in just a few genes that regulate _____ are often responsible for the evolution of new anatomical features in a population. (a) fossil formation (b) vestigial structures (c) development (d) biogeography (e) adaptation

CRITICAL THINKING Suzanne L. Collins and Joseph T. Collins/Photo Researchers, Inc.

1. EVOLUTION LINK. The use of model organisms such as the mouse for research and biomedical testing of human diseases is based on the assumption that all organisms share a common ancestor. On what evidence is this assumption based? 2. EVOLUTION LINK. What adaptations must an animal possess to swim in the ocean? Why are such genetically different organisms as porpoises, which are mammals, and sharks, which are fish, so similar in form? 3. EVOLUTION LINK. The human fetus grows a coat of fine hair (the lanugo) that is shed before or shortly after birth. Fetuses of chimpanzees and other primates also grow coats of hair, but they are not shed. Explain these observations based on what you have learned in this chapter. 4. EVOLUTION LINK. Charles Darwin once said, “It is not the strongest of the species that survive, nor the most intelligent, but the one most responsive to change.” Explain what he meant. 5. EVOLUTION LINK. Write short paragraphs explaining each of the following statements: a. Natural selection chooses from among the individuals in a population those most suited to current environmental conditions. It does not guarantee survival under future conditions. b. Individuals do not evolve, but populations do. c. The organisms that exist today do so because their ancestors had traits that allowed them and their offspring to thrive. d. At the molecular level, evolution can take place by the replacement of one nucleotide by another. e. Evolution is said to have occurred within a population when measurable genetic changes are detected. 6. EVOLUTION LINK. Although most salamanders have four legs, a few species that live in shallow water lack hind limbs and have extremely tiny forelimbs (see photograph). Develop a hypothesis to

The narrow-striped dwarf siren (Pseudobranchus striatus axanthus) is an aquatic salamander that resembles an eel. It is native to Florida.

explain how limbless salamanders came about according to Darwin’s mechanism of evolution by natural selection. How could you test your hypothesis? 7. ANALYZING DATA. Which of the primates in Figure 18-18 is the most distantly related to humans? Explain your answer. 8. SCIENCE, TECHNOLOGY, AND SOCIETY. Farmers often face a predicament, known as the pesticide treadmill, in which the cost of applying pesticides increases because the pesticides have to be applied more frequently or in larger doses, while their effectiveness decreases. Based on what you have learned in this chapter, offer an explanation for why the pesticides lose their effectiveness. Additional questions are available in CengageNOW at www.cengage.com/ login.

Evolutionary Change in Populations

G. I. Bernard/Animals Animals

19

Genetic variation in snail shells. Shown are the shell patterns and colors in a single snail species (Cepaea nemoralis), native to Scotland. Variation in shell color may have adaptive value in these snails because some colors predominate in cooler environments whereas other colors are more common in warmer habitats.

KEY CONCEPTS

A

19.1 The genotype, phenotype, and allele frequencies

ing differential survival and reproduction, individuals themselves do not

of a population’s gene pool (all alleles of all loci) can be calculated.

evolve during their lifetimes. Evolutionary change, which includes modi-

19.2 The Hardy–Weinberg principle predicts allele and

one generation to the next. Although Darwin recognized that evolution

genotype frequencies for a population that is not evolving.

19.3 Natural selection, as well as nonrandom mating, mutation, genetic drift, and gene flow are forces that drive changes in a population’s gene pool over successive generations (microevolution). Modes of natural selection include stabilizing selection, directional selection, and disruptive selection.

19.4 Populations typically exhibit genetic variation, which may include genetic polymorphism, balanced polymorphism, or geographic variation.

s you learned in Chapter 18, evolution occurs in populations, not individuals. Although natural selection acts on individuals, caus-

fications in structure, physiology, ecology, and behavior, is inherited from occurs in populations, he did not understand how the attributes of organisms are passed to successive generations. One of the most significant advances in biology since Darwin’s time has been the demonstration of the genetic basis of evolution. Recall from Chapter 18 that a population consists of all individuals of the same species that live in a particular place at the same time. Individuals within a population vary in many recognizable characters. A population of snails, for example, may vary in shell size, weight, or color (see photograph). Some of this variation is due to heredity, and some is due to environment (nonheritable variation), such as the individual differences observed in the pink color of flamingos, which is partly attributable to differences in diet. Biologists study variation in a particular character (characteristic) by taking measurements of that character in a population. By comparing the character in parents and offspring, it is possible to estimate the amount of observed variation that is genetic, as represented by the number, frequency, and kinds of alleles in a population. (Recall from

Chapter 11 that an allele is one of two or more alternate forms of a

Phenotype

Number

Phenotype Frequency

gene. Alleles occupy corresponding positions, or loci, on homolo-

Dominant Recessive Total

910 90 1000

0.91 0.09 1.00

gous chromosomes.) This chapter will help you develop an understanding of the importance of genetic variation as the raw material for evolution and of the basic concepts of population genetics, the study of genetic variability within a population and of the evolutionary forces that act on it. Population genetics represents an extension of Gregor Mendel’s principles of inheritance (see Chapter 11). You will learn how to distinguish genetic equilibrium from evolutionary change and to assess the roles of the five factors responsible for evolutionary change: nonrandom mating, mutation, genetic drift, gene flow, and natural selection.

In this example, the dominant phenotype is the sum of two genotypes, AA and Aa, so the number 910 is obtained by adding 490 + 420. An allele frequency is the proportion of a specific allele (that is, of A or a) in a particular population. As mentioned earlier, each individual, being diploid, has two alleles at each genetic locus. Because we started with a population of 1000 individuals, we must account for a total of 2000 alleles. The 490 AA individuals have 980 A alleles, whereas the 420 Aa individuals have 420 A alleles, making a total of 1400 A alleles in the population. The total number of a alleles in the population is 420 + 90 + 90 = 600. Now it is easy to calculate allele frequencies:

19.1 GENOTYPE, PHENOTYPE, AND ALLELE FREQUENCIES ■ ■ LEARNING OBJECTIVES 1 2

Define what is meant by a population’s gene pool. Distinguish among genotype, phenotype, and allele frequencies.

Each population possesses a gene pool, which includes all the alleles for all the loci present in the population. Because diploid organisms have a maximum of two different alleles at each genetic locus, a single individual typically has only a small fraction of the alleles present in a population’s gene pool. The genetic variation that is evident among individuals in a given population indicates that each individual has a different subset of the alleles in the gene pool. The evolution of populations is best understood in terms of genotype, phenotype, and allele frequencies. Suppose, for example, that all 1000 individuals of a hypothetical population have their genotypes tested, with the following results: Genotype AA Aa aa Total

Number

Genotype Frequency

490 420 90 1000

0.49 0.42 0.09 1.00

Each genotype frequency is the proportion of a particular genotype in the population. Genotype frequency is usually expressed as a decimal fraction, and the sum of all genotype frequencies is 1.0 (somewhat like probabilities, which were discussed in Chapter 11). For example, the genotype frequency for the Aa genotype is 420 ÷ 1000 = 0.42. A phenotype frequency is the proportion of a particular phenotype in the population. If each genotype corresponds to a specific phenotype, then the phenotype and genotype frequencies are the same. If allele A is dominant over allele a, however, the phenotype frequencies in our hypothetical population would be the following:

Allele

Number

Allele Frequency

A a Total

1400 600 2000

0.7 0.3 1.0

Review ■ ■

■

Does the term gene pool apply to individuals, populations, or both? Can the frequencies of all genotypes in a population be determined directly with respect to a locus that has only two alleles, one dominant and the other recessive? In a human population of 1000, 840 are tongue rollers (TT or Tt), and 160 are not tongue rollers (tt). What is the frequency of the dominant allele (T) in the population?

19.2 THE HARDY–WEINBERG PRINCIPLE ■ ■ LEARNING OBJECTIVES 3 4

Discuss the significance of the Hardy–Weinberg principle as it relates to evolution and list the five conditions required for genetic equilibrium. Use the Hardy–Weinberg principle to solve problems involving populations.

In the example just discussed, we observe that only 90 of the 1000 individuals in the population exhibit the recessive phenotype characteristic of the genotype aa. The remaining 910 individuals exhibit the dominant phenotype and are either AA or Aa. You might assume that after many generations, genetic recombination during sexual reproduction would cause the dominant allele to become more common in the population. You might also assume that the recessive allele would eventually disappear altogether. These were common assumptions of many biologists early in the 20th century. However, these assumptions are incorrect because the frequencies of alleles and genotypes do not change from generation to generation unless influenced by outside factors (discussed later in the chapter).

p2 Frequency of AA

+

2pq Frequency of Aa

+

q2 Frequency of aa

=

1 All individuals in the population

We always begin Hardy–Weinberg calculations by determining the frequency of the homozygous recessive genotype. From the fact that we had 90 homozygous recessive individuals in our population of 1000, we infer that the frequency of the aa genotype, q2,

Genotypes

AA

Aa

aa

Frequency of genotypes in population

0.49

0.42 (0.21 + 0.21)

0.09

Frequency of alleles in gametes

A = 0.49 + 0.21 = 0.7

a = 0.21 + 0.09 = 0.3

(a) Genotype and allele frequencies. The figure illustrates how to calculate frequencies of the alleles A and a in the gametes produced by each genotype.

Allele frequencies in female gametes

+ Allele frequencies in male gametes

A population whose allele and genotype frequencies do not change from generation to generation is said to be at genetic equilibrium. Such a population, with no net change in allele or genotype frequencies over time, is not undergoing evolutionary change. A population that is at genetic equilibrium is not evolving with respect to the locus being studied. However, if allele frequencies change over successive generations, evolution is occurring. The explanation for the stability of successive generations in populations at genetic equilibrium was provided independently by Godfrey Hardy, an English mathematician, and Wilhelm Weinberg, a German physician, in 1908. They pointed out that the expected frequencies of various genotypes in a population can be described mathematically. The resulting Hardy–Weinberg principle shows that if the population is large, the process of inheritance does not by itself cause changes in allele frequencies. It also explains why dominant alleles are not necessarily more common than recessive ones. The Hardy–Weinberg principle represents an ideal situation that seldom occurs in the natural world. However, it is useful because it provides a model to help us understand the real world. Knowledge of the Hardy–Weinberg principle is essential to understanding the mechanisms of evolutionary change in sexually reproducing populations. We now expand our original example to illustrate the Hardy– Weinberg principle. Keep in mind as we go through these calculations that in most cases we know only the phenotype frequencies. When alleles are dominant and recessive, it is usually impossible to visually distinguish heterozygous individuals from homozygous dominant individuals. The Hardy–Weinberg principle lets us use phenotype frequencies to calculate the expected genotype frequencies and allele frequencies, assuming we have a clear understanding of the genetic basis for the character under study. As mentioned earlier, the frequency of either allele, A or a, is represented by a number that ranges from 0 to 1. An allele that is totally absent from the population has a frequency of zero. If all the alleles of a given locus are the same in the population, then the frequency of that allele is 1. Because only two alleles, A and a, exist at the locus in our example, the sum of their frequencies must equal 1. If we let p represent the frequency of the dominant (A) allele in the population, and q the frequency of the recessive (a) allele, then we can summarize their relationship with a simple binomial equation, p + q = 1. When we know the value of either p or q, we can calculate the value of the other: p = 1 − q, and q = 1 − p. Squaring both sides of p + q = 1 results in (p + q)2 = 1. This equation can be expanded to describe the relationship of the allele frequencies to the genotypes in the population. When it is expanded, we obtain the frequency of the offspring genotypes:

A p = 0.7

a q = 0.3

A

a

p = 0.7

q = 0.3

AA

Aa

p2 = 0.7 × 0.7 = 0.49

pq = 0.7 × 0.3 = 0.21

Aa

aa

pq = 0.7 × 0.3 = 0.21

q2 = 0.3 × 0.3 = 0.09

(b) Segregation of alleles and random fertilization. When eggs and sperm containing A or a alleles unite randomly, the frequency of each of the possible genotypes (AA, Aa, aa) among the offspring is calculated by multiplying the frequencies of the alleles A and a in eggs and sperm.

FIGURE 19-1 Animated The Hardy–Weinberg principle

is 90/1000, or 0.09. Because q2 equals 0.09, q (the frequency of the recessive a allele) is equal to the square root of 0.09, or 0.3. From the relationship between p and q, we conclude that the frequency of the dominant A allele, p, equals 1 − q = 1 − 0.3 = 0.7. Given this information, we can calculate the frequency of homozygous dominant (AA) individuals: p2 = 0.7 × 0.7 = 0.49 (FIG. 19-1). The expected frequency of heterozygous individuals (Aa) would be 2pq = 2 × 0.7 × 0.3 = 0.42. Thus, approximately 490 individuals are expected to be homozygous dominant, and 420 are expected to be heterozygous. Note that the sum of homozygous dominant and heterozygous individuals equals 910, the number of individuals showing the dominant phenotype with which we started. Any population in which the distribution of genotypes conforms to the relation p2 + 2pq + q2 = 1, whatever the absolute values for p and q may be, is at genetic equilibrium. The Hardy– Weinberg principle allows biologists to calculate allele frequencies in a given population if we know the genotype frequencies, and vice versa. These values provide a basis of comparison with a

population’s allele or genotype frequencies in succeeding generations. During that time, if the allele or genotype frequencies deviate from the values predicted by the Hardy–Weinberg principle, then the population is evolving.

Genetic equilibrium occurs if certain conditions are met

blood group is of interest to population geneticists because the alleles for the MN blood group, usually designated M and N, are codominant (genotype MM produces antigen M only, genotype NN produces antigen N only, and the heterozygous genotype MN produces both antigens). This allows population geneticists to directly observe all three possible genotype frequencies and compare them with calculated frequencies. The following data are typical of the MN blood group in people in the United States:

The Hardy–Weinberg principle of genetic equilibrium tells us what to expect when a sexually reproducing population is not evolving. The relative proportions of alleles and genotypes in successive generations will always be the same, provided the following five conditions are met: 1. Random mating. In unrestricted random mating, each individual in a population has an equal chance of mating with any individual of the opposite sex. In our example, the individuals represented by the genotypes AA, Aa, and aa must mate with one another at random and must not select their mates on the basis of genotype or any other factors that result in nonrandom mating. 2. No net mutations. There must be no mutations that convert A into a, or vice versa. That is, the frequencies of A and a in the population must not change because of mutations. 3. Large population size. Allele frequencies in a small population are more likely to change by random fluctuations (that is, by genetic drift, which is discussed later) than are allele frequencies in a large population. 4. No migration. There can be no exchange of alleles with other populations that might have different allele frequencies. In other words, there can be no migration of individuals into or out of a population.1 5. No natural selection. If natural selection is occurring, certain phenotypes (and their corresponding genotypes) are favored over others. These more successful genotypes are said to have greater fitness, which is the relative ability to make a genetic contribution to subsequent generations. Consequently, the allele frequencies will change, and the population will evolve.

Human MN blood groups are a valuable illustration of the Hardy–Weinberg principle Humans have dozens of antigens on the surfaces of their blood cells. (An antigen is a molecule, usually a protein or carbohydrate, that is recognized as foreign by cells of another organism’s immune system.) One group of antigens, designated the MN blood group, stimulates the production of antibodies when injected into rabbits or guinea pigs. However, humans do not produce antibodies for M and N, so the MN blood group is not medically important, for example, when giving blood transfusions. (Recall the discussion of the medically important ABO alleles in Chapter 11.) The MN 1 Note that evolutionary biologists use the term migration, not in its ordinary sense of periodic or seasonal movement of individuals from one location to another but in reference to a movement of individuals that results in a transfer of alleles from one population to another.

Genotype

Observed

MM MN NN Total

320 480 200 1000

Because 1000 diploid individuals are in the sample, there are a total of 2000 alleles. The frequency of M alleles in the population = p = (2 × 320 + 480) ÷ 2000 = 0.56. The frequency of N alleles in the population = q = (2 × 200 + 480) ÷ 2000 = 0.44. As a quick check, the sum of the frequencies should equal 1. Does it? If this population is in genetic equilibrium, then the expected MM genotype frequency = p2 = (0.56)2 = 0.31. The expected MN genotype frequency = 2pq = 2 × 0.56 × 0.44 = 0.49. The expected NN genotype frequency = q2 = (0.44)2 = 0.19. As a quick check, the sum of the three genotype frequencies should equal 1. Does it? You can use the calculated genotype frequencies to determine how many individuals in a population of 1000 should have the expected genotype frequencies. By comparing the expected numbers with the actual results observed, you see how closely the population is to genetic equilibrium. Simply multiply each genotype frequency by 1000: Genotype MM MN NN Total

Observed

Expected

320 480 200 1000

313.6 492.8 193.6 1000.0

The expected numbers closely match the observed numbers, indicating that the MN blood groups in the human population are almost at genetic equilibrium. This is not surprising because the lack of medical significance suggests that the MN characteristic is not subject to natural selection and that it does not produce a visible phenotype that might affect random mating.

Review ■

■

■

In a population at genetic equilibrium, the frequency of the homozygous recessive genotype (tt) is 0.16. What are the allele frequencies of T and t, and what are the expected frequencies of the TT and Tt genotypes? In a population at genetic equilibrium, the frequency of the dominant phenotype is 0.96. What are the frequencies of the dominant (A) and recessive (a) alleles, and what are the expected frequencies of the AA, Aa, and aa genotypes? The genotype frequencies of a population are determined to be 0.6 BB, 0.0 Bb, and 0.4 bb. Is it likely that this population meets all the conditions required for genetic equilibrium?

■ ■ LEARNING OBJECTIVES 5 Define microevolution. 6 Discuss how each of the following microevolutionary forces alters allele

7

frequencies in populations: nonrandom mating, mutation, genetic drift, gene flow, and natural selection. Distinguish among stabilizing selection, directional selection, and disruptive selection.

Evolution represents a departure from the Hardy–Weinberg principle of genetic equilibrium. The degree of departure between the observed allele or genotype frequencies and those expected by the Hardy–Weinberg principle indicates the amount of evolutionary change. This type of evolution—generation-to-generation changes in allele or genotype frequencies within a population—is sometimes referred to as microevolution because it often involves relatively small or minor changes, usually over a few generations. Changes in the allele frequencies of a population result from five microevolutionary processes: nonrandom mating, mutation, genetic drift, gene flow, and natural selection. These microevolutionary processes are the opposite of the conditions that must be met if a population is in genetic equilibrium. When one or more of these processes acts on a population, allele or genotype frequencies change from one generation to the next.

Nonrandom mating changes genotype frequencies When individuals select mates on the basis of phenotype (thereby selecting the corresponding genotype), they bring about evolutionary change in the population. Two examples of nonrandom mating are inbreeding and assortative mating. In many populations, individuals mate more often with close neighbors than with more distant members of the population. As a result, neighbors tend to be more closely related—that is, genetically similar—to one another. The mating of genetically similar individuals who are more closely related than if they had been chosen at random from the entire population is known as inbreeding. Although inbreeding does not change the overall allele frequency, the frequency of homozygous genotypes increases with each successive generation of inbreeding. The most extreme example of inbreeding is self-fertilization, which is particularly common in certain plants. Inbreeding does not appear to be detrimental in some populations, but in others it causes inbreeding depression, in which inbred individuals have lower fitness than those not inbred. Fitness is usually measured as the average number of surviving offspring of one genotype compared to the average number of surviving offspring of competing genotypes. Inbreeding depression, as evidenced by fertility declines and high juvenile mortality, is thought to be caused by the expression of harmful recessive alleles as homozygosity increases with inbreeding. Several studies in the 1990s provided direct evidence of the deleterious consequences of inbreeding in nature. For example,

white-footed mice (Peromyscus leucopus) were taken from a field and used to develop both inbred and non-inbred populations in the laboratory. When these laboratory-bred populations were returned to nature, their survivorship was estimated from release–recapture data. The non-inbred mice had a statistically significant higher rate of survival (FIG. 19-2). It is not known why the inbred mice had a lower survival rate. Some possibilities include higher disease susceptibility, poorer ability to evade predators, less ability to find food, and less ability to win fights with other white-footed mice. Assortative mating, in which individuals select mates by their phenotypes, is another example of nonrandom mating. For example, biologists selected two phenotypes—high bristle number and low bristle number—in a fruit fly (Drosophila melanogaster) population. Although the researchers made no effort to control mating, they observed that the flies preferentially mated with those of similar phenotypes. Females with high bristle number tended to mate with males with high bristle number, and females with low bristle number tended to mate with males with low bristle number. Such

KEY EXPERIMENT QUESTION: Does inbreeding affect survival? HYPOTHESIS: Non-inbred white-footed mice (Peromyscus leucopus) will have a survival advantage over inbred mice in a natural environment. EXPERIMENT: Field-captured mice were used to establish inbred and non-inbred laboratory populations. The mice were marked and then released back into the field population. The population was then sampled by capturing mice, noting the number of inbred and non-inbred mice, and re-releasing the mice. The x-axis represents the 10-week period during which the samples were collected. Values on the y-axis are the estimated proportion of mice that survived from one week to the next. For example, the value of 0.2 in week 3 for the inbred mice means that 20% of the inbred mice that were alive at the beginning of the week survived through that week. 1.4 1.2 Weekly survival

19.3 MICROEVOLUTION

Non-inbred mice

1.0 0.8 0.6

Inbred mice

0.4 0.2 1

2

3

4 5 6 7 8 Weeks after release

9

10

11

RESULTS AND CONCLUSION: Non-inbred mice (red) had a consistently higher survival rate than inbred mice (blue). Thus, inbreeding had a negative effect on survival in this species in nature. Adapted with permission from Jiménez, J. A., et al., “An Experimental Study of Inbreeding Depression in a Natural Habitat,” Science, Vol. 266, Oct. 14, 1994. Copyright © 1994 American Association for the Advancement of Science.

FIGURE 19-2 Survival of inbred and non-inbred mice

selection of mates with the same phenotype is known as positive assortative mating (as opposed to the less common phenomenon, negative assortative mating, in which mates with opposite phenotypes are selected). Positive assortative mating is practiced in many human societies, in which men and women tend to marry individuals like themselves in such characteristics as height or intelligence. Like inbreeding, assortative mating usually increases homozygosity at the expense of heterozygosity in the population and does not change the overall allele frequencies in the population. However, assortative mating changes genotype frequencies only at the loci involved in mate choice, whereas inbreeding affects genotype frequencies in the entire genome.

Mutation increases variation within a population Variation is introduced into a population through mutation, which is an unpredictable change in deoxyribonucleic acid (DNA). As discussed in Chapter 13, mutations, which are the source of all new alleles, result from (1) a change in the nucleotide base pairs of a gene, (2) a rearrangement of genes within chromosomes so that their interactions produce different effects, or (3) a change in chromosome structure. Mutations occur unpredictably and spontaneously. A particular locus may have a DNA sequence that causes certain types of mutations to occur more frequently than others. The rate of mutation appears relatively constant for a particular locus but may vary by several orders of magnitude among genes within a single species and among different species. Not all mutations pass from one generation to the next. Those occurring in somatic (body) cells are not inherited. When an individual with such a mutation dies, the mutation is lost. Some mutations, however, occur in reproductive cells. These mutations may or may not overtly affect the offspring because most of the DNA in a cell is “silent” and does not code for specific polypeptides or proteins that are responsible for physical characteristics. Even if a mutation occurs in the DNA that codes for a polypeptide, it may still have little effect on the structure or function of that polypeptide (we discuss such neutral variation later in the chapter). However, when a polypeptide is sufficiently altered to change its function, the mutation is usually harmful. By acting against seriously abnormal phenotypes, natural selection eliminates or reduces to low frequencies the most harmful mutations. Mutations with small phenotypic effects, even if slightly harmful, have a better chance of being incorporated into the population, where at some later time, under different environmental conditions, they may produce phenotypes that are useful or adaptive. The “needs” of a population do not determine what mutations will occur. Consider a population living in an increasingly dry environment. A mutation producing a new allele that helps an individual adapt to dry conditions is no more likely to occur than one for adapting to wet conditions or one with no relationship to the changing environment. The production of new mutations simply increases the genetic variability that is acted on by natural selection and, therefore, increases the potential for new adaptations. Mutation by itself causes small deviations in allele frequencies from those predicted by the Hardy–Weinberg principle. Although

allele frequencies may be changed by mutation, these changes are typically several orders of magnitude smaller than changes caused by other evolutionary forces, such as genetic drift. Mutation is usually negligible as an evolutionary force, but it is essential to the evolutionary process because it is the ultimate source of genetic variation.

In genetic drift, random events change allele frequencies The size of a population has important effects on allele frequencies because random events, or chance, tend to cause changes of relatively greater magnitude in a small population. If a population consists of only a few individuals, an allele present at a low frequency in the population could be completely lost by chance. Such an event would be unlikely in a large population. For example, consider two populations, one with 10,000 individuals and one with 10 individuals. If an uncommon allele occurs at a frequency of 10%, or 0.1, in both populations, then 1900 individuals in the large population have the allele.2 That same frequency, 0.1, in the smaller population means that only about 2 individuals have the allele.3 From this exercise, it is easy to see that there is a greater likelihood of losing the rare allele from the smaller population than from the larger one. Predators, for example, might happen to kill one or two individuals possessing the uncommon allele in the smaller population purely by chance, so these individuals would leave no offspring. The production of random evolutionary changes in small breeding populations is known as genetic drift. Genetic drift results in changes in allele frequencies in a population from one generation to another. One allele may be eliminated from the population purely by chance, regardless of whether that allele is beneficial, harmful, or of no particular advantage or disadvantage. Thus, genetic drift decreases genetic variation within a population, although it tends to increase genetic differences among different populations. When bottlenecks occur, genetic drift becomes a major evolutionary force Because of fluctuations in the environment, such as depletion in food supply or an outbreak of disease, a population may rapidly and markedly decrease from time to time. The population is said to go through a bottleneck during which genetic drift can occur in the small population of survivors. As the population again increases in size, many allele frequencies may be quite different from those in the population preceding the decline. Scientists hypothesize that genetic variation in the cheetah was considerably reduced by a bottleneck at the end of the last Ice Age, some 10,000 years ago (see Fig. 52-9). Cheetahs nearly became extinct, perhaps from overhunting by humans. The few surviving cheetahs had greatly reduced genetic variability, and as a result, the cheetah population today is so genetically uniform that unrelated cheetahs can accept skin grafts from one another. (Normally, only identical twins accept skin grafts so readily.) 2pq + q2 = 2(0.9)(0.1) + (0.1)2 = 0.18 + 0.01= 0.19; 0.19 × 10,000 = 1900

2

3

0.19 × 10 = 1.9

The Finnish and Icelandic populations, as well as certain Amish populations, are being studied by researchers in the quest to identify genetic contributions to a large number of diseases. The task is simplified in these studies because confounding factors such as variability in the rest of the genome and differences in such environmental factors as nutrition, access to medical care, and exposure to pollutants are relatively controlled.

The founder effect occurs when a few “founders” establish a new colony When one or a few individuals from a large population establish, or found, a colony (as when a few birds separate from the rest of the flock and fly to a new area), they bring with them only a small fraction of the genetic variation present in the original population. As a result, the only alleles among their descendants will be those of the colonizers. Typically, the allele frequencies in the newly founded population are quite different from those of the parent population. The genetic drift that results when a small number of individuals from a large population found a new colony is called the founder effect. The Finnish people may illustrate the founder effect (FIG. 19-3). Geneticists who sampled DNA from Finns and from the European population at large found that Finns exhibit considerably less genetic variation than other Europeans. This evidence supports the hypothesis that Finns are descended from a small group of people who settled about 4000 years ago in the area that is now Finland and, because of the geography, remained separate from other European societies for centuries. The subpopulation in eastern Finland is especially homogeneous because it existed in relative isolation after being established by only several hundred founders in the 1500s. The founder effect and population bottlenecks have also apparently affected the genetic composition of the population of Iceland. Iceland’s 275,000 citizens are descended from a small group of Norse and Celtic people who settled that island in the ninth century. Isolation and population bottlenecks due to disease and natural disasters have contributed to the relative genetic homogeneity of the Icelandic people. The founder effect can be of medical importance. For example, by chance one of the approximately 200 founders of the Amish population of Pennsylvania carried a recessive allele that, when homozygous, is responsible for a form of dwarfism, Ellis–van Creveld syndrome. Although this allele is rare in the general population (frequency about 0.001), today it is relatively common in the Amish population (frequency about 0.07).

Gene flow generally increases variation within a population Individuals of a species tend to be distributed in local populations that are genetically isolated to some degree from other populations. For example, the bullfrogs of one pond form a population separated from those in an adjacent pond. Some exchanges occur by migration between ponds, but the frogs in one pond are much more likely to mate with those in the same pond. Because each population is isolated to some extent from other populations, each has distinct genetic traits and gene pools. The migration of breeding individuals between populations causes a corresponding movement of alleles, or gene flow, that has significant evolutionary consequences. As alleles flow from one population to another, they usually increase the amount of genetic variability within the recipient population. If sufficient gene flow occurs between two populations, they become more similar genetically. Because gene flow reduces the amount of variation between two populations, it tends to counteract the effects of natural selection and genetic drift, both of which often cause populations to become increasingly distinct. If migration by members of a population is considerable and if populations differ in their allele frequencies, then significant genetic changes occur in local populations. For example, by 10,000 years ago modern humans occupied almost all of Earth’s major land areas except a few islands. Because the population density was low in most locations, the small, isolated human populations underwent random genetic drift and natural selection. More recently (during the past 300 years or so), major migrations have increased gene flow, significantly altering allele frequencies within previously isolated human populations.

Yann Arthus Bertrand/Corbis

Natural selection changes allele frequencies in a way that increases adaptation

FIGURE 19-3 Finns and the founder effect The Finnish people are thought to have descended from a small founding population that remained separate from the rest of Europe for centuries.

Natural selection is the mechanism of evolution first proposed by Darwin in which members of a population that are more successfully adapted to the environment have greater fitness—that is, they are more likely to survive and reproduce (see Chapter 18). Over successive generations, the proportion of favorable alleles increases in the population. In contrast with other microevolutionary processes (nonrandom mating, mutation, genetic drift, and gene flow), natural selection leads to adaptive evolutionary change. Natural selection not only explains why organisms are well adapted to the environments in which they live but also helps account for the remarkable diversity of life. Natural selection enables populations to change, thereby adapting to different environments and different ways of life.

Natural selection results in the differential reproduction of individuals with different traits, or phenotypes (and therefore different genotypes), in response to the environment. Natural selection preserves individuals with favorable phenotypes and eliminates those with unfavorable phenotypes. Individuals that survive and produce fertile offspring have a selective advantage. The mechanism of natural selection does not develop a “perfect” organism. Rather, it weeds out those individuals whose phenotypes are less adapted to environmental challenges, while allowing better-adapted individuals to survive and pass their alleles to their offspring. By reducing the frequency of alleles that result in the expression of less favorable traits, the probability is increased that favorable alleles responsible for an adaptation will come together in the offspring. Natural selection operates on an organism’s phenotype Natural selection does not act directly on an organism’s genotype. Instead, it acts on the phenotype, which is, at least in part, an expression of the genotype. The phenotype represents an interaction between the environment and all the alleles in the organism’s genotype. It is rare that alleles of a single locus determine the phenotype, as Mendel originally observed in garden peas. Much more common is the interaction of alleles of several loci for the expression of a single phenotype. Many plant and animal characteristics are under this type of polygenic control (see Chapter 11). When characters are under polygenic control (as is human height), a range of phenotypes occurs, with most of the population located in the median range and fewer at either extreme. This

KEY POINT

is a normal distribution or standard bell curve (FIG. 19-4a; see also Fig. 11-20). Three kinds of selection cause changes in the normal distribution of phenotypes in a population: stabilizing, directional, and disruptive selection. Although we consider each process separately, in nature their influences generally overlap. Stabilizing Selection The process of natural selection associated with a population well adapted to its environment is known as stabilizing selection. Most populations are probably influenced by stabilizing forces most of the time. Stabilizing selection selects against phenotypic extremes. In other words, individuals with average, or intermediate, phenotypes are favored. Because stabilizing selection tends to decrease variation by favoring individuals near the mean of the normal distribution at the expense of those at either extreme, the bell curve narrows (FIG. 19-4b). Although stabilizing selection decreases the amount of variation in a population, variation is rarely eliminated by this process because other microevolutionary processes act against a decrease in variation. For example, mutation is slowly but continually adding to the genetic variation within a population. One of the most widely studied cases of stabilizing selection involves human birth weight, which is under polygenic control and is also influenced by environmental factors. Extensive data from hospitals have shown that infants born with intermediate weights are most likely to survive (FIG. 19-5). Infants at either extreme (too small or too large) have higher rates of mortality. When newborn infants are too small, their body systems are immature, and when they are too large, they have difficult deliveries because they cannot pass as easily through the cervix and vagina. Stabilizing selec-

Stabilizing, directional, and disruptive selection can change the distribution of phenotypes in a population.

Stabilizing selection

Directional selection

Disruptive selection

Number of individuals

No selection

Phenotype

(a) A character that is under polygenic control (in this example, wing colors in a hypothetical population of beetles) exhibits a normal distribution of phenotypes in the absence of selection.

(b) As a result of stabilizing selection, which trims off extreme phenotypes, variation about the mean is reduced.

(c) Directional selection shifts the curve in one direction, changing the average value of the character.

FIGURE 19-4 Animated Modes of selection The blue screen represents the distribution of individuals by phenotype (in this example, color variation) in the original population. The purple screen represents the distribution by phenotype in the evolved population. The arrows represent the pressure of natural selection on the phenotypes.

(d) Disruptive selection, which trims off intermediate phenotypes, results in two or more peaks.

100

15

30

10

10 5

5 2 1

2

3

4

5

6

7

8

Body weight in pounds

9 10 11

Percentage of infant death — log scale (red)

Percentage of infant population (blue)

20

TABLE 19-1

Population Changes in Geospiza fortis before and after the 1976–1977 Drought

Character Weight (g) Wing length (mm) Tarsus (leg, just above the foot) length (mm) Bill length (mm) Bill depth (mm) Bill width (mm)

Average before Average after Drought (634)* Drought (135)* Difference 16.06 67.88

17.13 68.87

+1.07 +0.99

19.08 10.63 9.21 8.58

19.29 10.95 9.70 8.83

+0.21 +0.32 +0.49 +0.25

From P. R. Grant and B. R. Grant, “Predicting Microevolutionary Responses to Directional Selection on Heritable Variation,” Evolution, Vol. 49, 1995. *Number of birds in sample.

FIGURE 19-5 Stabilizing selection The blue curve shows the distribution of birth weights in a sample of 13,730 infants. The red curve shows mortality (death) at each birth weight. Infants with very low or very high birth weights have higher death rates than infants of average weight. The optimum birth weight, that is, the one with the lowest mortality, is close to the average birth weight (about 7 lb). (Adapted from Cavalli-Sforza, L. L., and W. F. Bodmer, The Genetics of Human Populations, W. H. Freeman and Company, San Francisco, 1971.)

Giuliano Gerra and Silvio Sommazzi

tion operates to reduce variability in birth weight so it is close to the weight with the minimum mortality rate. tional selection may favor phenotypes at one of the extremes of the normal distribution (FIG. 19-4c). Over successive generations, one phenotype gradually replaces another. So, for example, if greater size is advantageous in a new environment, larger individuals will become increasingly common in the population. However, directional selection occurs only if alleles favored under the new circumstances are already present in the population. Darwin’s Galápagos finches provide an excellent example of directional selection. Since 1973, Peter Grant and Rosemary Grant of Princeton University have studied the Galápagos finches. The Grants did a meticulous analysis of finch eating habits and beak sizes on Isla Daphne Major during three extended droughts (1977–1978, 1980, and 1982), one of which was followed by an extremely wet El Niño event (1983). During the droughts, the number of insects and small seeds declined, and large, heavy seeds became the finches’ primary food source. Many finches died during this time, and most of the survivors were larger birds whose beaks were larger and deeper. In a few generations, these larger birds became more common in the population (TABLE 19-1). After the El Niño event, however, smaller seeds became the primary food source, and smaller finches with averagesized beaks were favored. In this example, natural selection is directional: during the drought, natural selection operated in favor of the larger phenotype; whereas after the wet period, selection occurred in the opposite direction, favoring the smaller phenotype. The guppy populations studied in Venezuela and Trinidad (see Chapter 18) are another example of directional selection. Directional selection by hunters has been observed in bighorn sheep (Ovis canadensis) in Canada. Sport hunters target large rams with rapidly growing horns for their trophy value (FIG. 19-6).

Harry Engels/Photo Researchers, Inc.

Directional Selection If an environment changes over time, direc-

FIGURE 19-6 Bighorn sheep This ram exhibits the massive horns that characterize this species.

Because these rams are typically killed prior to reaching their full reproductive potential, they contribute less to the gene pool; the result has been a 25% decline in average ram weight and horn length over a period of 30 years. Paradoxically, the rams that would

normally have the greatest fitness because of their size and ability to use their horns in competition with other males for mates become less fit when these attributes cause them to be singled out by hunters.

variation. The sexual process allows the variability introduced by mutation to be combined in new ways, which may be expressed as new phenotypes.

Disruptive Selection Sometimes extreme changes in the environ-

Genetic polymorphism can be studied in several ways

ment may favor two or more different phenotypes at the expense of the mean. That is, more than one phenotype may be favored in the new environment. Disruptive selection is a special type of directional selection in which there is a trend in several directions rather than just one (FIG. 19-4d). It results in a divergence, or splitting apart, of distinct groups of individuals within a population. Disruptive selection, which is relatively rare, selects against the average, or intermediate, phenotype. Limited food supply during a severe drought caused a population of finches on another island in the Galápagos to experience disruptive selection. The finch population initially exhibited a variety of beak sizes and shapes. Because the only foods available on this island during the drought were wood-boring insects and seeds from cactus fruits, natural selection favored birds with beaks suitable for obtaining these types of food. Finches with longer beaks survived because they could open cactus fruits, and those with wider beaks survived because they could strip off tree bark to expose insects. However, finches with intermediate beaks could not use either food source efficiently and had a lower survival rate. Natural selection induces change in the types and frequencies of alleles in populations only if there is pre-existing inherited variation. Genetic variation is the raw material for evolutionary change because it provides the diversity on which natural selection acts. Without genetic variation, evolution cannot occur. In the next section we explore the genetic basis for variation that is acted on by natural selection.

Review ■

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Which microevolutionary force leads to adaptive changes in allele frequencies? Why is mutation important to evolution if it is the microevolutionary force that generally has the smallest effect on allele frequencies? Which microevolutionary forces are most associated with an increase in variation within a population? among populations? Which microevolutionary force typically changes genotype frequencies without changing allele frequencies? Explain.

19.4 GENETIC VARIATION IN POPULATIONS ■ ■ LEARNING OBJECTIVE 8

Describe the nature and extent of genetic variation, including genetic polymorphism, balanced polymorphism, neutral variation, and geographic variation.

Populations contain abundant genetic variation that was originally introduced by mutation. Sexual reproduction, with its associated crossing-over, independent assortment of chromosomes during meiosis, and random union of gametes, also contributes to genetic

Genetic polymorphism, which is genetic variation among individuals in a population, is extensive in populations, although many of the genetic variants are present at low frequencies. Much of genetic polymorphism is not evident because it does not produce distinct phenotypes. One way biologists estimate the total amount of genetic polymorphism in populations is by comparing the different forms of a particular protein. Each form consists of a slightly different amino acid sequence that is coded for by a different allele. For example, tissue extracts containing a particular enzyme may be analyzed for different individuals by gel electrophoresis. In gel electrophoresis, the enzymes are placed in slots on an agarose gel and an electric current is applied, causing each enzyme to migrate across the gel (see Fig. 15-8). Slight variations in amino acid sequences in the different forms of a particular enzyme cause each to migrate at a different rate, which can be detected using special stains or radioactive labels. Depending on the species studied, the percentage of enzymes that are polymorphic can range from about 15% to as much as 70%. In general, polymorphism of enzyme-coding loci tends to be greater in plants than in animals. Determining the sequence of nucleotides in DNA from individuals in a population provides a direct estimate of genetic polymorphism that is sensitive enough to detect alleles that differ by only one nucleotide (single nucleotide polymorphisms, SNPs). (DNA sequencing is shown in Figures 15-10 and 15-11.) DNA sequencing of specific alleles in an increasing number of organisms, including humans, indicates that SNPs are common in most populations. Recent studies have revealed that genetic changes in which segments of DNA have been gained or lost (compared to a reference genome) are common in model organisms such as mice and Drosophila, as well as humans. These copy number variations, or CNVs, can involve DNA segments that range from as few as 500 to 1000 base pairs up to about two million base pairs. It is currently estimated that the average human may have a dozen or more CNVs in their genome. In most cases, there does not appear to be an associated phenotype, or the phenotype is benign, such as an effect on eye color. However, as more sophisticated technologies have been developed to support ongoing research, evidence is accumulating that links specific CNVs to increased risk for certain medically important conditions, such as autism, Alzheimer’s disease, and susceptibility to HIV infection.

Balanced polymorphism exists for long periods Balanced polymorphism is a special type of genetic polymorphism in which two or more alleles persist in a population over many generations as a result of natural selection. Heterozygote advantage and frequency-dependent selection are mechanisms that preserve balanced polymorphism.

Genetic variation may be maintained by heterozygote advantage We have seen that natural selection often eliminates unfavorable alleles from a population, whereas favorable alleles are retained. However, natural selection sometimes helps maintain genetic diversity. In some cases natural selection may even maintain a population’s alleles that are unfavorable in the homozygous state. This happens, for example, when the heterozygote Aa has a higher degree of fitness than either homozygote AA or aa. This phenomenon, known as heterozygote advantage, is demonstrated in humans by the selective advantage of heterozygous carriers of the sickle cell allele. The mutant allele (HbS) for sickle cell anemia produces an altered hemoglobin that deforms or sickles the red blood cells, making them more likely to form dangerous blockages in capillaries and to be destroyed in the liver, spleen, or bone marrow (discussed in Chapter 16). People who are homozygous for the sickle cell allele (HbSHbS) usually die at an early age if medical treatment is not available. Heterozygous individuals carry alleles for both normal (HbA) and sickle cell hemoglobin. The heterozygous condition (HbAHbS) makes a person more resistant to a type of severe malaria (caused by the parasite Plasmodium falciparum) than people who are homozygous for the normal hemoglobin allele (HbAHbA). In a heterozygous individual, each allele produces its own specific kind of hemoglobin, and the red blood cells contain the two kinds in roughly equivalent amounts. Such cells sickle much less readily than cells containing only the HbS allele. They are more resistant to infection by the malaria-causing parasite, which lives in red blood cells, than are the red blood cells containing only normal hemoglobin. Where malaria is a problem, each of the two types of homozygous individuals is at a disadvantage. Those homozygous for the sickle cell allele are likely to die of sickle cell anemia, whereas those homozygous for the normal allele may die of malaria. The heterozygote is therefore more fit than either homozygote. In parts of Af-

Europe

Genetic variation may be maintained by frequency-dependent selection Thus far in our discussion of natural selection, we have assumed that the fitness of particular phenotypes (and their corresponding genotypes) is independent of their frequency in the population. However, in cases of frequency-dependent selection, the fitness of a particular phenotype depends on how frequently it appears in the population. Often a phenotype has a greater selective value when rare than when common in the population. Such phenotypes lose their selective advantage as they become more common. Frequency-dependent selection often acts to maintain genetic variation in populations of prey species. In this case, the predator catches and consumes the more common phenotype but may ignore the rarer phenotypes. Consequently, the less common phenotype produces more offspring and therefore makes a greater relative contribution to the next generation. Frequency-dependent selection is demonstrated in scaleeating fish (cichlids of the species Perissodus microlepsis) from Lake

Europe

Asia

Africa

Atlantic Ocean

rica, the Middle East, and southern Asia where falciparum malaria is prevalent, heterozygous individuals survive in greater numbers than either homozygote (FIG. 19-7). The HbS allele is maintained at a high frequency in the population, even though the homozygous recessive condition is almost always lethal. What happens to the frequency of HbS alleles in Africans and others who possess it when they migrate to the United States and other countries with few cases of malaria? As might be expected, the frequency of the HbS allele gradually declines in such populations, possibly because it confers a selective disadvantage by causing sickle cell anemia in homozygous individuals but no longer confers a selective advantage by preventing malaria in heterozygous individuals. The HbS allele never disappears from the population, however, because it is “hidden” from selection in heterozygous individuals and because it is reintroduced into the population by gene flow from the African population.

Asia

Africa

Indian Ocean

Atlantic Ocean

Frequency of Hb s allele 1–10% 10–20%

Australia

(a) The geographic distribution of falciparum malaria (green).

Indian Ocean Australia

(b) The distribution of the sickle cell anemia allele (red and orange).

FIGURE 19-7 Heterozygote advantage Over large geographic areas, the distribution of individuals heterozygous for the sickle cell (Hbs) allele is similar to the distribution of falciparum malaria. This evidence for greater fitness of heterozygous individuals in malarial regions supports the hypothesis of heterozygote advantage. (Adapted from Allison, A. C., “Protection Afforded by Sickle-Cell Traits against Subtertian Malarial Infection,” British Medical Journal, Vol. 1, 1954.)

Tanganyika in Africa (FIG. 19-8). The scale-eating fish, which obtain food by biting scales off other fish, have either left-pointing or right-pointing mouths. A single locus with two alleles determines this characteristic; the allele for right-pointing mouth is dominant

KEY EXPERIMENT QUESTION: Can evidence for frequency-dependent selection be found in natural populations? HYPOTHESIS: Equal proportions of scale-eating cichlids with right-pointing mouths vs. left-pointing mouths in a population are maintained by frequency-dependent selection. Cichlid with right-pointing mouth attacks prey on its left flank.

over the allele for left-pointing mouth. These fish attack their prey from behind and from a single direction, depending on mouth morphology. Those with left-pointing mouths always attack the right flanks of their prey, whereas those with right-pointing mouths always attack the left flanks. The prey species are more successful at evading attacks from the more common form of scale-eating fish. For example, if the cichlids with right-pointing mouths are more common than those with left-pointing mouths, the prey are attacked more often on their left flanks. They therefore become more wary against such attacks, conferring a selective advantage to the less common cichlids with left-pointing mouths. The cichlids with left-pointing mouths would be more successful at obtaining food and would therefore have more offspring. Over time, the frequency of fish with leftpointing mouths would increase in the population, until their abundance causes frequency-dependent selection to work against them and confer an advantage on the now less common fish with rightpointing mouths. Thus, frequency-dependent selection maintains both populations of fish at approximately equal numbers.

Neutral variation may give no selective advantage or disadvantage Cichlid with left-pointing mouth attacks prey on its right flank.

Frequency of cichlid individuals with left-pointing mouths

EXPERIMENT: The relative frequencies of scale-eating cichlids with left-pointing and right-pointing mouths in Lake Tanganyika in Africa were monitored over a 10-year period. 1.0

0.5

0

1982

1984

1986

1988

1990

Sample years

RESULTS AND CONCLUSION: Relative frequencies of leftpointing and right-pointing cichlids varied slightly from year to year, but remained close to 0.5. This approximate 1:1 ratio is consistent with frequency-dependent selection because prey fish become especially wary to attacks by the cichlid type that is currently most common; therefore either type is at a selective disadvantage if its frequency rises above 0.5.

FIGURE 19-8 Frequency-dependent selection in scale-eating cichlids (Adapted with permission from Hori, M., “Frequency-Dependent Natural Selection in the Handedness of Scale-Eating Cichlid Fish,” Science, Vol. 260, April 9, 1993. Copyright © 1993 American Association for the Advancement of Science.)

Some of the genetic variation observed in a population may confer no apparent selective advantage or disadvantage in a particular environment. For example, random changes in DNA that do not alter protein structure usually do not affect the phenotype. Variation that does not alter the ability of an individual to survive and reproduce and is, therefore, not adaptive is called neutral variation. The extent of neutral variation in organisms is difficult to determine. It is relatively easy to demonstrate that an allele is beneficial or harmful if its effect is observable. But the variation in alleles that involves only slight differences in the proteins they code for may or may not be neutral. These alleles may be influencing the organism in subtle ways that are difficult to measure or assess. Also, an allele that is neutral in one environment may be beneficial or harmful in another.

Populations in different geographic areas often exhibit genetic adaptations to local environments In addition to the genetic variation among individuals within a population, genetic differences often exist among different populations within the same species, a phenomenon known as geographic variation. One type of geographic variation is a cline, which is a gradual change in a species’s phenotype and genotype frequencies through a series of geographically separate populations as a result of an environmental gradient. A cline exhibits variation in the expression of such attributes as color, size, shape, physiology, or behavior. Clines are common among species with continuous ranges over large geographic areas. For example, the body sizes of many widely distributed birds and mammals increase gradually as the latitude increases, presumably because larger animals are better able to withstand the colder temperatures of winter. The common yarrow (Achillea millefolium), a wildflower that grows in a variety of North American habitats from lowlands to

Mean height of yarrow plants (cm)

100

50

0

Groveland Mather

Aspen Yosemite Tenaya Tuolumne Valley Creek Lake Meadows

Big Timberline Conway Horn Summit Lake

Lee Vining

Elevation (m)

4000 3000 2000 1000 0 Yarrow (Achillea millefolium)

Sierra Nevada Range

Great Basin Plateau

FIGURE 19-9 Clinal variation in yarrow (Achillea millefolium) (After J. Clausen, D. D. Keck, and W. M. Hiesey, “Experimental Studies on the Nature of Species: III. Environmental Responses of Climatic Races of Achillea,” Carnegie Institute Washington Publication, Vol. 58, 1948.)

mountain highlands, exhibits clinal variation in height in response to different climates at different elevations. Although substantial variation exists among individuals within each population, individuals in populations at higher elevations are, on average, shorter than those at lower elevations. The genetic basis of these clinal differences was experimentally demonstrated in a set of classical experiments in which series of populations from different geographic areas were grown in the same environment (FIG. 19-9). Despite being exposed to identical environmental conditions, each experi-

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mental population exhibited the traits characteristic of the elevation from which it was collected.

Review ■ ■

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How does the sickle cell allele illustrate heterozygote advantage? How does frequency-dependent selection affect genetic variation within a population over time? How can researchers test the hypothesis that clinal variation among populations of a particular species has a genetic basis?

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

19.1 (page 412) 1 Define what is meant by a population’s gene pool. All the individuals that live in a particular place at the same time make up a population. Each population has a gene pool, which includes all the alleles for all the loci present in the population. ■ Population genetics is the study of genetic variability within a population and of the forces that act on it. 2 Distinguish among genotype, phenotype, and allele frequencies. ■ A genotype frequency is the proportion of a particular genotype in the population. A phenotype frequency is the proportion of a particular phenotype in the population. An allele frequency is the proportion of a specific allele of a given genetic locus in the population. ■

19.2 (page 412) 3 Discuss the significance of the Hardy–Weinberg principle as it relates to evolution and list the five conditions required for genetic equilibrium.

The Hardy–Weinberg principle states that allele and genotype frequencies do not change from generation to generation (no evolution is occurring) in a population at genetic equilibrium. ■ The Hardy–Weinberg principle applies only if mating is random in the population, there are no net mutations that change the allele frequencies, the population is large, individuals do not migrate between populations, and natural selection does not occur. 4 Use the Hardy–Weinberg principle to solve problems involving populations. ■ In the Hardy–Weinberg equation, p = the frequency of the dominant allele, q = the frequency of the recessive allele: p + q = 1. The genotype frequencies of a population are described by the ■ relationship p2 + 2pq + q2 = 1, where p2 is the frequency of the homozygous dominant genotype, 2pq is the frequency of the heterozygous genotype, and q2 is the frequency of the homozygous recessive genotype. ■

Interact with the Hardy–Weinberg principle by clicking on the figure in CengageNOW.

5 Define microevolution. Microevolution is a change in allele or genotype frequencies within a population over successive generations. 6 Discuss how each of the following microevolutionary forces alters allele frequencies in populations: nonrandom mating, mutation, genetic drift, gene flow, and natural selection. ■ In nonrandom mating, individuals select mates on the basis of phenotype, indirectly selecting the corresponding genotype(s). Inbreeding is the mating of genetically similar individuals that are more closely related than if they had been chosen at random from the entire population. Inbreeding in some populations causes inbreeding depression, in which inbred individuals have lower fitness (relative ability to make a genetic contribution to the next generation) than non-inbred individuals. In assortative mating individuals select mates by their phenotypes. Both inbreeding and assortative mating increase the frequency of homozygous genotypes. ■ Mutations, unpredictable changes in DNA, are the source of new alleles. Mutations increase the genetic variability acted on by natural selection. ■ Genetic drift is a random change in the allele frequencies of a small population. Genetic drift decreases genetic variation within a population, and the changes caused by genetic drift are usually not adaptive. A sudden decrease in population size caused by adverse environmental factors is known as a bottleneck. The founder effect is genetic drift that occurs when a small population colonizes a new area. ■ Gene flow, a movement of alleles caused by the migration of individuals between populations, causes changes in allele frequencies. ■ Natural selection causes changes in allele frequencies that lead to adaptation. Natural selection operates on an organism’s phenotype, but it changes the genetic composition of a population in a favorable direction for a particular environment. 7 Distinguish among stabilizing selection, directional selection, and disruptive selection. ■

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Stabilizing selection favors the mean at the expense of phenotypic extremes. Directional selection favors one phenotypic extreme over another, causing a shift in the phenotypic mean. Disruptive selection favors two or more phenotypic extremes.

Harry Engels, Photo Researchers, Inc.

19.3 (page 415)

Watch natural selection in action by clicking on the figure in CengageNOW.

19.4 (page 420) 8 Describe the nature and extent of genetic variation, including genetic polymorphism, balanced polymorphism, neutral variation, and geographic variation. ■ Genetic polymorphism is the presence of genetic variation within a population. ■ Balanced polymorphism is a special type of genetic polymorphism in which two or more alleles persist in a population over many generations as a result of natural selection. ■ Heterozygote advantage occurs when the heterozygote exhibits greater fitness than either homozygote. In frequency-dependent selection, a genotype’s selective value varies with its frequency of occurrence. ■ Neutral variation is genetic variation that confers no detectable selective advantage. ■ Geographic variation is genetic variation that exists among different populations within the same species. A cline is a gradual change in a species’s phenotype and genotype frequencies through a series of geographically separate populations.

T E S T YO U R U N D E R S TA N D I N G 1. The genetic description of an individual is its genotype, whereas the genetic description of a population is its (a) phenotype (b) gene pool (c) genetic drift (d) founder effect (e) changes in allele frequencies 2. In a diploid species, each individual possesses (a) one allele for each locus (b) two alleles for each locus (c) three or more alleles for each locus (d) all the alleles found in the gene pool (e) half of the alleles found in the gene pool 3. The MN blood group is of interest to population geneticists because (a) people with genotype MN cannot receive blood transfusions from either MM or NN people (b) the MM, MN, and NN genotype frequencies can be observed directly and compared with calculated expected frequencies (c) the M allele is dominant to the N allele (d) people with the MN genotype exhibit frequencydependent selection (e) people with the MN genotype exhibit heterozygote advantage 4. If all copies of a given locus have the same allele throughout the population, then the allele frequency is (a) 0 (b) 0.1 (c) 0.5 (d) 1.0 (e) 10.0

5. If a population’s allele and genotype frequencies remain constant from generation to generation, (a) the population is undergoing evolutionary change (b) the population is said to be at genetic equilibrium (c) microevolution has taken place (d) directional selection is occurring, but only for a few generations (e) genetic drift is a significant evolutionary force 6. Analysis of DNA sequences for a particular locus among individuals within a population provides biologists with one way to estimate (a) genetic drift (b) genetic polymorphism (c) gene flow (d) heterozygote advantage (e) frequency-dependent selection 7. The continued presence of the allele that causes sickle cell anemia in areas where falciparum malaria is prevalent demonstrates which of the following phenomena? (a) inbreeding depression (b) frequency-dependent selection (c) heterozygote advantage (d) genetic drift (e) a genetic bottleneck 8. According to the Hardy–Weinberg principle, (a) allele frequencies are not dependent on dominance or recessiveness but remain essentially unchanged from generation to generation (b) the sum of allele frequencies for a given locus is always greater than 1 (c) if a

locus has only one allele, the frequency of that allele is zero (d) allele frequencies change from generation to generation (e) the process of inheritance, by itself, causes changes in allele frequencies 9. The Hardy–Weinberg principle may be applicable if (a) population size is small (b) migration occurs only at the beginning of the breeding season (c) mutations occur at a constant rate (d) matings occur exclusively between individuals of the same genotype (e) natural selection does not occur

by preserving the traits acquired during an individual’s lifetime (d) natural selection acts on a population’s genetic variability, which arises through mutation (e) natural selection may result in changes in allele frequencies in a population 12. The distribution of phenotypes shown in the figure is most consistent with (a) no selection (b) stabilizing selection (c) directional selection (d) disruptive selection (e) genetic equilibrium

10. Mutation (a) leads to adaptive evolutionary change (b) adds to the genetic variation of a population (c) is the result of genetic drift (d) almost always benefits the organism (e) a and b 11. Which of the following is not true of natural selection? (a) natural selection acts to preserve favorable traits and eliminate unfavorable traits (b) the offspring of individuals that are better adapted to the environment will make up a larger proportion of the next generation (c) natural selection directs the course of evolution

CRITICAL THINKING 1. EVOLUTION LINK. Why are mutations almost always neutral or harmful? If this is true, why are mutations nevertheless essential to evolution? 2. EVOLUTION LINK. Explain this apparent paradox: scientists discuss evolution in terms of genotype fitness (the selective advantage that a particular genotype confers on an individual), yet natural selection acts on an organism’s phenotype. 3. EVOLUTION LINK. Why is it easier for researchers to study the genetic contribution to disease in the populations of Finland or Iceland, as opposed to those of the United States? 4. EVOLUTION LINK. Evolution is sometimes characterized as “survival of the fittest.” Is this consistent with an evolutionary biologist’s definitions of fitness and natural selection? Is this a good way to think about evolution, or a poor one? 5. ANALYZING DATA. Recall that the allele for right-pointing mouth in cichlids (A) is dominant over that for left-pointing mouth (a). Use the Hardy–Weinberg principle to calculate the allele frequencies that would correspond to equal frequencies of the two phenotypes. Which allele has the higher frequency? Why? How are these scale-eating fish an example of balanced polymorphism? 6. ANALYZING DATA. The recessive allele that causes Ellis–van Creveld syndrome when homozygous has a frequency of about 0.07 in the Amish population of Pennsylvania, although its fre-

quency is only about 0.001 in the general population. How many persons out of 1000 in the Amish population would be expected to have the disease? How many out of a million in the general population? 7. ANALYZING DATA. You study males in populations of a certain species of minnows in a series of lakes at different latitudes. You find that they exhibit clinal variation in average weight at maturity and hypothesize that the weight differences are due to genetic factors. You predict that the average weights at maturity of representatives of each population reared in aquaria will differ in ways that are consistent with the differences you observed among the wild populations. However, when you conduct the experiment, you find no differences among the population averages. What is your conclusion? 8. SCIENCE, TECHNOLOGY, AND SOCIETY. Automated techniques are being developed that will provide new information on the extent of genetic polymorphism among humans, particularly with respect to single nucleotide polymorphisms (SNPs) and copy number variations (CNVs). In what ways might this information prove useful? In what ways might it present challenges? Additional questions are available in CengageNOW at www.cengage.com/ login.

20

Speciation and Macroevolution

© iStockphoto.com/Mariko Yuki

Blue-footed booby. Blue-footed boobies are large sea birds that nest on the Galápagos Islands and several other arid islands in the eastern Pacific Ocean. Male and female pairs are monogamous and typically nest on black lava. The largest blue-footed booby breeding population is found in the Galápagos Islands.

KEY CONCEPTS 20.1 According to the biological species concept, a species consists of individuals that can successfully interbreed with one another but not with individuals from other species. 20.2 The evolution of different species begins with repro-

A

Brazilian rain forest contains thousands of species of insects, amphibians, reptiles, birds, and mammals. The Great Barrier Reef off the

coast of Australia has thousands of species of sponges, corals, mollusks, crustaceans, sea stars, and fishes. The various environments on Earth abound in rich assemblages of species. Scientists have currently named about 1.8 million species. We do not know exactly how many species exist today, but many

ductive isolation, in which two populations are no longer able to interbreed successfully.

biologists estimate the number to be on the order of 4 million to

20.3 In allopatric speciation, populations diverge into

100 million! Often this diversity has been portrayed as many twig tips

different species due to geographic isolation, or physical separation. In sympatric speciation, populations become reproductively isolated from one another despite living in the same geographic area.

on a “tree of life.” As one follows the twigs from the tips toward the main

20.4 Speciation may require millions of years but some-

body of the tree, they form connections that represent common ancestors that are now extinct. Indeed, more than 99.9% of all species that ever existed are now extinct. Tracing the branches of the tree of life to larger

times occurs much more quickly.

and larger connections eventually leads to a single trunk that represents

20.5 The evolution of species and higher taxa is known

the few simple unicellular organisms that evolved early in Earth’s history

as macroevolution.

and became the ancestors of all species living today.

Thus, life today is the product of 3 billion to 4 billion years of evolution. How did all these species diversify throughout Earth’s history? Darwin’s studies led him to conclude that the Galápagos Islands were the birthplace of many new species (see photograph). Our understanding of how species evolve has advanced significantly since Darwin published On the Origin of Species. In this chapter we consider the reproductive barriers that isolate species from one another and the possible evolutionary mechanisms that explain how the millions of species that live today or lived in the past originated from ancestral species. We then examine the rate of evolutionary change and the evolution of higher taxonomic categories (species, genus, family, order, class,

20.1 WHAT IS A SPECIES? ■ ■ LEARNING OBJECTIVE 1

Describe the biological species concept and list two potential problems with the concept.

The concept of distinct kinds of organisms, known as species (from Latin, meaning “kind”), is not new. However, there are several different definitions of species, and every definition has some sort of limitation. Linnaeus was the 18th-century biologist who founded modern taxonomy—the science of naming, describing, and classifying organisms. He classified plants and other organisms into separate species based on their visible structural differences, such as feathers or number of flower parts. This method, known as the morphological species concept, is still used to help characterize species, but structural differences alone are not adequate to explain what constitutes a species.

The biological species concept is based on reproductive isolation Population genetics did much to clarify the concept of species. According to the biological species concept, first expressed by evolutionary biologist Ernst Mayr in 1942, a species consists of one or more populations whose members interbreed in nature to produce fertile offspring and do not interbreed with—that is, are reproductively isolated from—members of different species. Gene flow occurs between individuals of a given species when they mate, but reproductive barriers restrict individuals of one species from interbreeding, or exchanging genes, with members of other species. As a result, reproductive barriers keep a given species genetically distinct from other species. An extension of the biological species concept holds that new species evolve when formerly interbreeding populations become reproductively isolated from one another. The biological species concept is currently the most widely accepted definition of species, but it has several shortcomings. One problem with the biological species concept is that it applies only to sexually reproducing organisms. Bacteria reproduce asexually,

Wim van Egmond/Visuals Unlimited, Inc.

and phylum), which is the focus of macroevolution.

150 μm

FIGURE 20-1 Bdelloid rotifer (Philodina roseola) Bdelloid rotifers are all females. Because they do not reproduce sexually, it is difficult to classify these small marine animals as species under the biological species concept.

although they may have considerable exchange of genetic material by means of horizontal gene transfer among bacterial individuals that are classified as different species. Thus, reproductive isolation is not a good criterion for defining species of bacteria. Small marine organisms known as bdelloid rotifers are an example of animals that reproduce asexually (FIG. 20-1). All bdelloid rotifers are females, and they produce offspring without needing males. These organisms and also extinct organisms must be classified based on their structural and biochemical characteristics; the biological species concept does not apply to them. Another potential problem with the biological species concept is that although individuals assigned to different species do not normally interbreed, they may sometimes successfully interbreed. For example, where their ranges overlap, some coyotes have been known to mate with wolves. As a result, some wolves carry coyote genes and vice versa. Yet, most biologists agree that wolves and coyotes are separate species.

The phylogenetic species concept defines species based on such evidence as molecular sequencing Some biologists prefer the phylogenetic species concept, also called the evolutionary species concept, in which a population is

declared a separate species if it has undergone evolution long enough for statistically significant differences in diagnostic traits to emerge. If a population has a unique diagnostic trait—a set of spines running down the back, for example—that can be demonstrated to be inherited, then this population is considered a separate species. This approach has the advantage of being testable by comparing gene sequences between two groups. However, many biologists do not want to abandon the biological species concept, in part because the phylogenetic species concept cannot be applied if the evolutionary history (as determined by DNA sequencing) of a taxonomic group has not been carefully studied; most groups have not been rigorously analyzed. Also, if the phylogenetic species concept were universally embraced, the number of named species would probably double. This increase would occur because many closely related populations that are classified as subspecies (a taxonomic subdivision of a species) under the biological species concept would fit the requirements of separate species under the phylogenetic species concept. Thus, the exact definition of a species remains fuzzy. Unless we clearly state otherwise, when we mention species in this text, we are referring to the biological species concept.

Review ■

What is a species, according to the biological species concept? What are some of the limitations of this definition?

20.2 REPRODUCTIVE ISOLATION ■ ■ LEARNING OBJECTIVE 2

Explain the significance of reproductive isolating mechanisms and distinguish among the various prezygotic and postzygotic barriers.

Various reproductive isolating mechanisms prevent interbreeding between two different species whose ranges (areas where each lives) overlap. These mechanisms preserve the genetic integrity of each species because gene flow between the two species is prevented.

Most species have two or more mechanisms that block a chance occurrence of individuals from two closely related species overcoming a single reproductive isolating mechanism. Although most mechanisms operate before mating or fertilization occurs (prezygotic), others work after fertilization has taken place (postzygotic).

Prezygotic barriers interfere with fertilization Prezygotic barriers are reproductive isolating mechanisms that prevent fertilization from taking place. Because male and female gametes never come into contact, an interspecific zygote—that is, a fertilized egg formed by the union of an egg from one species and a sperm from another species—is never produced. Prezygotic barriers include temporal isolation, habitat isolation, behavioral isolation, mechanical isolation, and gametic isolation. Sometimes genetic exchange between two groups is prevented because they reproduce at different times of the day, season, or year. Such examples demonstrate temporal isolation. For example, two very similar species of fruit flies, Drosophila pseudoobscura and D. persimilis, have ranges that overlap to a great extent, but they do not interbreed. Drosophila pseudoobscura is sexually active only in the afternoon; and D. persimilis, only in the morning. Similarly, two frog species have overlapping ranges in eastern Canada and the United States. The wood frog (Rana sylvatica) usually mates in late March or early April, when the water temperature is about 7.2°C (45°F), whereas the northern leopard frog (R. pipiens) usually mates in mid-April, when the water temperature is 12.8°C (55°F) (FIG. 20-2). Although two closely related species may be found in the same geographic area, they usually live and breed in different habitats in that area. This provides habitat isolation between the two species. For example, the five species of small birds known as flycatchers are nearly identical in appearance and have overlapping ranges in the eastern part of North America. They exhibit habitat isolation because during the breeding season, each species stays in a particular habitat within its range, so potential mates from different species do not meet. The least flycatcher (Empidonax minimus)

(a) The wood frog (Rana sylvatica) mates in early spring, often before the ice has completely melted in the ponds.

(b) The leopard frog (Rana pipiens) typically mates a few weeks later.

FIGURE 20-2 Animated Temporal isolation in wood and leopard frogs

Leopard frog

Mating activity

Rod Planck/Photo Researchers, Inc.

L. & D. Klein/Photo Researchers, Inc.

Wood frog

March 1

April 1

May 1

(c) Graph of peak mating activity in wood and leopard frogs. In nature, wood and leopard frogs do not interbreed.

R. Brown/VIREO

FIGURE 20-3 Behavioral isolation in bowerbirds Each bowerbird species has highly specialized courtship patterns that prevent its mating with another species. The male satin bowerbird (Ptilonorhynchus violacens) constructs an enclosed place, or bower, of twigs to attract a female. (The bower is the dark “tunnel” on the left.) Note the white flowers and blue decorations, including human-made objects such as bottle caps, that he has arranged at the entrance to his bower.

Sometimes members of different species court and even attempt copulation, but the incompatible structures of their genital organs prevent successful mating. Structural differences that inhibit mating between species produce mechanical isolation. For example, many flowering plant species have physical differences in their flower parts that help them maintain their reproductive isolation from one another. In such plants, the flower parts are adapted for specific insect pollinators. Two species of sage, for example, have overlapping ranges in southern California. Black sage (Salvia mellifera), which is pollinated by small bees, has a floral structure different from that of white sage (S. apiana), which is pollinated by large carpenter bees (FIG. 20-4). Interestingly, black sage and white sage are also prevented from mating by a temporal barrier: black sage flowers in early spring, and white sage flowers in late spring and early summer. Presumably, mechanical isolation prevents insects from cross-pollinating the two species should they happen to flower at the same time. If mating takes place between two species, their gametes may still not combine. Molecular and chemical differences between species cause gametic isolation, in which the egg and sperm of different species are incompatible. In aquatic animals that release their eggs and sperm into the surrounding water simultaneously, interspecific fertilization is extremely rare. The surface of the egg contains specific proteins that bind only to complementary molecules on the surface of sperm cells of the same species (see Chapter 51). A similar type of molecular recognition often occurs between pollen grains and the stigma (receptive surface of the

frequents open woods, farms, and orchards; whereas the acadian flycatcher (E. virescens) is found in deciduous forests, particularly in beech trees, and swampy woods. The alder flycatcher (E. alnorum) prefers wet thickets of alders, the yellow-bellied flycatcher (E. flaviventris) nests in conifer woods, and the willow flycatcher (E. traillii) frequents brushy pastures and willow thickets. Many animal species exchange a distinctive series of signals before mating. Such courtship behaviors illustrate behavioral isolation (also known as sexual isolation). Bowerbirds, for example, exhibit species-specific courtship patterns. The male satin bowerbird of Australia constructs an elaborate bower of twigs, adding decorative blue parrot feathers and white flowers at the entrance (FIG. 20-3). When a female approaches the bower, the male dances around her, holding a particularly eye-catching decoration in his beak. While dancing, he puffs his feathers, extends his wings, and sings a courtship song that consists of a variety of sounds, including loud buzzes and hoots. These specific courtship behaviors keep closely related bird species reproductively isolated from the satin bowerbird. If a male and female of two different bowerbird species begin courtship, it stops when one member does not recognize or respond to the signals of the other. Another example of behav(a) The petal of the black sage functions (b) The larger landing platform and longer stamens ioral isolation involves the wood as a landing platform for small bees. Large of white sage allow pollination by larger California frogs and northern leopard frogs bees cannot fit on this platform. carpenter bees (a different species). If smaller bees just discussed as an example of land on white sage, their bodies do not brush temporal isolation. Males of these against the stamens. (In the figure, the upper part of two species have very specific vothe white sage flower has been removed.) calizations to attract females of their species for breeding. These FIGURE 20-4 Mechanical isolation in black sage and white sage vocalizations reinforce each speThe differences in their floral structures allow black sage and white sage to be pollinated by different insects. Because the two species exploit different pollinators, they cannot interbreed. cies’s reproductive isolation.

of a cross between two sunflower species was 80%. In other words, 80% of the F2 generation were defective in some way and could not reproduce successfully. Hybrid breakdown also occurs in the F3 and later generations. TABLE 20-1 summarizes the various prezygotic and postzygotic barriers that prevent interbreeding between two species.

female part of the flower) so that pollen does not germinate on the stigma of a different plant species.

Postzygotic barriers prevent gene flow when fertilization occurs Fertilization sometimes occurs between gametes of two closely related species despite the existence of prezygotic barriers. When this happens, postzygotic barriers that increase the likelihood of reproductive failure come into play. Generally, the embryo of an interspecific hybrid spontaneously aborts. Development, the orderly sequence of events that occurs as an organism grows and matures, is a complex process requiring the precise interaction and coordination of many genes. Apparently, the genes from parents belonging to different species do not interact properly in regulating the mechanisms for normal development. In this case, reproductive isolation occurs by hybrid inviability. For example, in crosses between different species of irises the embryos die before reaching maturity. Similarly, nearly all the hybrids die in the embryonic stage when the eggs of a bullfrog are fertilized artificially with sperm from a leopard frog. If an interspecific hybrid does live, it may not reproduce, for several reasons. Hybrid animals may exhibit courtship behaviors incompatible with those of either parental species, and as a result they will not mate. More often, hybrid sterility occurs when problems during meiosis cause the gametes of an interspecific hybrid to be abnormal. Hybrid sterility is particularly common if the two parental species have different chromosome numbers. For example, a mule is the offspring of a female horse (2n = 64) and a male donkey (2n = 62) (FIG. 20-5). This type of union almost always results in sterile offspring (2n = 63) because synapsis (the pairing of homologous chromosomes during meiosis) and segregation of chromosomes cannot occur properly. Occasionally, a mating between two F1 hybrids produces a second hybrid generation (F2). The F2 hybrid may exhibit hybrid breakdown, the inability of a hybrid to reproduce because of some defect. For example, hybrid breakdown in the F2 generation

Biologists are discovering genes responsible for reproductive isolating mechanisms Progress has been made in identifying some of the genes involved in reproductive isolation. For example, scientists have determined the genetic basis for prezygotic isolation in species of abalone, large mollusks found along the Pacific coast of North America. In abalone, the fertilization of eggs by sperm requires lysin, a sperm protein that attaches to a lysin receptor protein located on the egg envelope. After attachment, the lysin produces a hole in the egg envelope that permits the sperm to penetrate the egg. Scientists cloned the lysin receptor gene and demonstrated that this gene varies among abalone species. Differences in the lysin receptor protein in various abalone species determine sperm compatibility with the egg. Sperm of one abalone species do not attach to a lysin receptor protein of an egg of a different abalone species. Other genes related to reproductive isolating mechanisms— loosely referred to as “speciation genes”—have been identified in a diverse group of organisms, such as Florida beach mice, cactus finches, fruit fly (Drosophila) species, and flycatcher birds in the South Pacific.

Review ■

■ ■

John Eastcott/Yva Momatiuk/Earth Scenes/Animals Animals

■

FIGURE 20-5 Hybrid sterility in mules Mules are interspecific hybrids formed by mating a female horse (left) with a male donkey (right). Although the mule (center) exhibits valuable characteristics of each of its parents, it is sterile.

What barriers prevent wood frogs and leopard frogs from interbreeding in nature? How is temporal isolation different from behavioral isolation? How is mechanical isolation different from gametic isolation? Of which postzygotic barrier is the mule an example?

20.3 SPECIATION ■ ■ LEARNING OBJECTIVES 3 Explain the mechanism of allopatric speciation and give an example. 4 Explain the mechanisms of sympatric speciation and give both plant and animal examples.

We are now ready to consider how entirely new species may arise from previously existing ones. The evolution of a new species is called speciation. The formation of two species from a single species occurs when a population becomes reproductively isolated from other populations of the species and the gene pools of the two separated populations begin to diverge in genetic composition. When a population is sufficiently different from its ancestral species that there is no genetic exchange between them, we consider speciation to have occurred. Speciation happens in two ways: allopatric speciation and sympatric speciation.

TABLE 20-1

Animated

Mechanism

Reproductive Isolating Mechanisms

How It Works

Example

Prezygotic barriers prevent fertilization Temporal isolation

Similar species reproduce at different times

Two species of fruit flies of the genus Drosophila live in the same geographic area, but one is sexually active only in the afternoon and the other only in the morning.

Habitat isolation

Similar species reproduce in different habitats

Several flycatcher species of the genus Empidonax live in the same geographic area but in different habitats; for example, the least flycatcher in open woods, farms, and orchards, and the acadian flycatcher in deciduous forests and swampy woods.

Behavioral isolation

Similar species have distinctive courtship behaviors

The male satin bowerbird of Australia constructs an elaborate bower, dances around the female, and sings a specific courtship song that keeps closely related bowerbird species reproductively isolated from the satin bowerbird.

Mechanical isolation

Similar species have structural differences in their reproductive organs

Black sage (genus Salvia) is pollinated by small bees, whereas white sage is pollinated by large carpenter bees. Because of differences in the flower structures, neither type of bee can successfully pollinate the other sage species.

Gametic isolation

Gametes of similar species are chemically incompatible

In aquatic animals such as sponges that release their eggs and sperm into the water simultaneously, interspecific fertilization rarely occurs because the egg surface is compatible only with the sperm surface of the same species.

Postzygotic barriers reduce viability or fertility of hybrid Hybrid inviability

Interspecific hybrid dies at early stage of embryonic development

In crosses between different species of irises (genus Iris), fertilization occurs but the embryos in the seeds die before reaching maturity.

Hybrid sterility

Interspecific hybrid survives to adulthood but is unable to reproduce successfully

A mule, the offspring of a female horse and a male donkey, has sterile offspring because synapsis and segregation of chromosomes do not occur properly during gamete formation.

Hybrid breakdown

Offspring of interspecific hybrid are unable to reproduce successfully

In a cross between two sunflower species, most of the F2 generation are defective in some way and cannot reproduce successfully. Hybrid breakdown also occurs in later generations.

up, leaving isolated pools. Presumably, each pool contained a small population of pupfish that gradually diverged from the common ancestral species by genetic drift and natural selection in response to habitat differences—the high temperatures, high salt concentrations, and low oxygen levels characteristic of desert springs. Today, there are 20 or so distinct species, subspecies, and populations in the area of Death Valley, California, and Ash Meadows, Nevada (FIG. 20-6). Many, such as the Devil’s Hole pupfish and the Owens pupfish, are restricted to one or two isolated springs. Allopatric speciation also occurs when a small population migrates or is dispersed (such as by a chance storm) and colonizes a new area away from the range of the original species. This colony is geographically isolated from its parental species, and the small microevolutionary changes that accumulate in the isolated gene pool over many generations may eventually be sufficient to form a new species. Because islands provide the geographic isolation required for allopatric speciation, they offer excellent opportunities to study this evolutionary mechanism. A few individuals of a few species probably colonized the Galápagos Islands and the Hawaiian Islands, for example. The hundreds of unique species presently found on each island presumably descended from these original colonizers (FIG. 20-7). Speciation is more likely to occur if the original isolated population is small. Recall from Chapter 19 that genetic drift, including the founder effect, is more consequential in small populations. Genetic drift tends to result in rapid changes in allele frequencies in a small, isolated population. The different selective pressures of the new environment to which the population is exposed further accentuate the divergence caused by genetic drift.

Long physical isolation and different selective pressures result in allopatric speciation

The Kaibab squirrel is an example of allopatric speciation in progress About 10,000 years ago, when the American Southwest was less arid, the ponderosa pine forests in the area supported a tree squir-

©Stone Nature Photography/Alamy

©Stone Nature Photography/Alamy

Speciation that occurs when one population becomes geographically separated from the rest of the species and subsequently evolves by natural selection and/or genetic drift is known as allopatric speciation (from the Greek allo, “different,” and patri, “native land”). Recall from Chapter 18 that natural selection occurs as individuals that possess favorable adaptations to the environment survive and become parents of the next generation. Genetic drift is a random change in allele frequency resulting from the effects of chance on the survival and reproductive success of individuals in a small breeding population (see Chapter 19). Both natural selection and genetic drift result in changes in allele frequencies in a population, but only in natural selection is the change in allele frequency adaptive. Allopatric speciation is the most common method of speciation and accounts for almost all evolution of new animal species. The geographic isolation required for allopatric speciation may occur in several ways. Earth’s surface is in a constant state of change. Such change includes rivers shifting their courses; glaciers migrating; mountain ranges forming; land bridges forming that separate previously united aquatic populations; and large lakes diminishing into several smaller, geographically separated pools. What may be an imposing geographic barrier to one species may be of no consequence to another. Birds and cattails, for example, do not become isolated when a lake subsides into smaller pools; birds easily fly from one pool to another, and cattails disperse their pollen and fruits by air currents. Fishes, in contrast, usually cannot cross the land barriers between the pools and so become reproductively isolated. In the Death Valley region of California and Nevada, large interconnected lakes formed during wetter climates of the last Ice Age. These lakes were populated by one or several species of pupfishes. Over time, the climate became drier, and the large lakes dried

(a) Devil‘s Hole pupfish (Cyprinodon diabolis), photographed at Devil‘s Hole, Ash Meadows National Wildlife Refuge, Nevada.

(b) Ash Meadows pupfish (Cyprinodon nevadensis ssp. mionectes), photographed at Point of Rocks Springs, Ash Meadows National Wildlife Refuge, Nevada.

FIGURE 20-6 Allopatric speciation of pupfishes (Cyprinodon) Shown are two pupfish species that evolved when larger lakes in southern Nevada dried up about 10,000 years ago, leaving behind small, isolated desert pools fed by springs. The pupfish’s short, stubby body is characteristic of fish that live in springs; fish that live in larger bodies of water are more streamlined.

FIGURE 20-7 Allopatric speciation of the Hawaiian goose (the nene) Nene (pronounced “nay-nay”; Branta sandvicensis) are geese originally found only on volcanic mountains on the geographically isolated islands of Hawaii and Maui, which are some 4200 km (2600 mi) from the nearest continent. Compared with those of other geese, the feet of Hawaiian geese are not completely webbed, their toenails are longer and stronger, and their foot pads are thicker; these adaptations enable Hawaiian geese to walk easily on lava flows. Nenes probably evolved from a small population of geese that originated in North America. Photographed in Hawaii Volcanoes National Park, Hawaii.

Porto Santo rabbits diverged rapidly from European rabbits

Allopatric speciation has the potential to occur quite rapidly. Early in the 15th century, a small population of rabbits was released on Porto Santo, a small island off the coast of Portugal. Because there were no other rabbits or competitors and no predators on the island, the rabbits thrived. By the 19th century, these rabbits were markedly different from their European ancestors. They were only half as large (weighing slightly more than 500 g, or 1.1 lb), with a different color pattern and a more nocturnal lifestyle. More significant, attempts to mate Porto Santo rabbits with mainland European rabbits failed. Many biologists concluded that within 400 years, an extremely brief period in evolutionary history, a new species of rabbit had evolved. Not all biologists agree that the Porto Santo rabbit is a new species. The objection stems from a more recent breeding experiment and is based on biologists’ lack of a consensus about the definition of a species. In the experiment, foster mothers of the wild Mediterranean rabbit raised newborn Porto Santo rabbits. When they reached adulthood, these Porto Santo rabbits mated successfully with Mediterranean rabbits to produce healthy, fertile offspring. To some biologists, this experiment clearly demonstrated that Porto Santo rabbits are not a separate species but a subspecies. These biologists cite the Porto Santo rabbits as an example of speciation in progress, much like the Kaibab squirrels just discussed. Many other biologists think the Porto Santo rabbit is a separate species because it does not interbreed with the other rabbits under natural conditions. They point out that the breeding experiment was successful only after the baby Porto Santo rabbits were raised under artificial conditions that probably modified their natural behavior.

Tom and Pat Leeson

rel with conspicuous tufts of hair sprouting from its ears. A small tree squirrel population living on the Kaibab Plateau of the Grand Canyon became geographically isolated when the climate changed, thus causing areas to the north, west, and east to become desert. Just a few miles to the south lived the rest of the squirrels, which we know as Abert squirrels, but the two groups were separated by the Grand Canyon. With changes over time in both its appearance and its ecology, the Kaibab squirrel is on its way to becoming a new species. During its many years of geographic isolation, the small population of Kaibab squirrels has diverged from the widely distributed Abert squirrels in several ways. Perhaps most evident are changes in fur color. The Kaibab squirrel now has a white tail and a gray belly, in contrast to the gray tail and white belly of the Abert squirrel (FIG. 20-8). Biologists think these striking changes arose in Kaibab squirrels as a result of genetic drift. Scientists have not reached a consensus about whether the two squirrel populations are separate (a) The Kaibab squirrel, with its white tail (b) The Abert squirrel, with its gray tail species, and most biologists today and gray belly, is found north of the Grand and white belly, is found south of the Canyon. Grand Canyon. consider the Kaibab squirrel (Sciurus aberti ssp. kaibabensis) and the FIGURE 20-8 Animated Allopatric speciation in progress Abert squirrel (Sciurus aberti) as

Kent and Donna Dannen

Victoria McCormick/Earth Scenes/Animals Animals

distinct subspecies. Because the Kaibab and Abert squirrels remain reproductively isolated from each other, it is assumed that they will continue to evolve into separate species.

Two populations diverge in the same physical location by sympatric speciation Although geographic isolation is an important factor in many cases of evolution, it is not an absolute requirement. In sympatric speciation (from the Greek sym, “together,” and patri, “native land”), a new species evolves within the same geographic region as the parental species. The divergence of two populations in the same geographic range occurs when reproductive isolating mechanisms evolve at the start of the speciation process. Sympatric speciation is especially common in plants. The role of sympatric speciation in animal evolution is probably much less important than allopatric speciation; until recently, sympatric speciation in animals has been difficult to demonstrate in nature. Sympatric speciation occurs in at least two ways: a change in ploidy (the number of chromosome sets making up an organism’s genome) and a change in ecology. We now examine each of these mechanisms. Allopolyploidy is an important mechanism of sympatric speciation in plants As a result of reproductive isolating mechanisms discussed earlier, the union of two gametes from different species rarely forms viable offspring; if offspring are produced, they are usually sterile. Synapsis cannot usually occur in interspecific hybrid offspring because not all the chromosomes are homologous. However, if the chromosome number doubles before meiosis, then homologous chromosomes can undergo synapsis. Although not common, this spontaneous doubling of chromosomes has been documented in a variety of plants and a few animals. It produces nuclei with multiple sets of chromosomes. Polyploidy, the possession of more than two sets of chromosomes, is a major factor in plant evolution. Reproductive isolation occurs in a single generation when a polyploid species with multiple sets of chromosomes arises from diploid parents. There are two kinds of polyploidy: autopolyploidy and allopolyploidy. An autopolyploid contains multiple sets of chromosomes from a single species, and an allopolyploid contains multiple sets of chromosomes from two or more species. We discuss only allopolyploidy because it is much more common in nature. Allopolyploidy occurs in conjunction with hybridization, which is sexual reproduction between individuals from closely related species. Allopolyploidy produces a fertile interspecific hybrid because the polyploid condition provides the homologous chromosome pairs necessary for synapsis during meiosis. As a result, gametes may be viable (FIG. 20-9). An allopolyploid—that is, an interspecific hybrid produced by allopolyploidy—reproduces with itself (self-fertilization) or with a similar individual. However, allopolyploids are reproductively isolated from both parents because their gametes have a different number of chromosomes from those of either parent. If a population of allopolyploids (that is, a new species) becomes established, selective pressures cause one of three outcomes. First, the new species may not compete successfully against species that are already established, so it becomes extinct. Second, the allopolyploid individuals may assume a new role in the environment and so coexist with both parental species. Third, the new species

KEY POINT

Changes in chromosome number have often led to speciation in plants. Species B 2n = 4

Species A 2n = 6 P generation

n=2

n=3 Gametes

Hybrid AB

F1 generation

No doubling of chromosome number

Doubling of chromosome number 2n = 10

Chromosomes either cannot pair or go through erratic meiosis.

No gametes or sterile gametes– no sexual reproduction is possible.

Pairing is now possible during meiosis.

n=5

Viable gametes; self-fertilization produces 2n = 10 offspring.

FIGURE 20-9 Sympatric speciation by allopolyploidy

in plants When two species (designated the P generation) successfully interbreed, the interspecific hybrid offspring (the F1 generation) are almost always sterile (bottom left). If the chromosomes double, proper synapsis and segregation of the chromosomes can occur and viable gametes may be produced (bottom right). (Unduplicated chromosomes are shown for clarity.)

may successfully compete with either or both of its parental species. If it has a combination of traits that confers greater fitness than one or both parental species for all or part of the original range of the parent(s), the hybrid species may replace the parent(s). Although allopolyploidy is extremely rare in animals, it is significant in the evolution of flowering plant species. As many as 80% of all flowering plant species are polyploids, and most of these are allopolyploids. Moreover, allopolyploidy provides a mechanism

for extremely rapid speciation. A single generation is all that is needed to form a new, reproductively isolated species. Allopolyploidy may explain the rapid appearance of many flowering plant species in the fossil record and their remarkable diversity today (at least 300,000 species). The kew primrose (Primula kewensis) is an example of sympatric speciation that was documented at the Royal Botanic Gardens at Kew, England, in 1898 (FIG. 20-10). Plant breeders developed P. kewensis as an interspecific hybrid of two primrose species, P. floribunda (2n = 18) and P. verticillata (2n = 18). Primula kewensis had a chromosome number of 18 but was sterile. Then, at three different times, it was reported to have spontaneously formed a fertile branch, which was an allopolyploid (2n = 36) that produced viable seeds of P. kewensis. The mechanism of sympatric speciation has been experimentally verified for many plant species. One example is a group of species, collectively called hemp nettles, that occur in temperate parts of Europe and Asia. One hemp nettle, Galeopsis tetrahit (2n = 32), is a naturally occurring allopolyploid hypothesized to have formed by the hybridization of two species, G. pubescens (2n = 16) and G. speciosa (2n = 16). This process occurred in nature but was experimentally reproduced. Galeopsis pubescens and G. speciosa were crossed to produce F1 hybrids, most of which were sterile. Nevertheless, both F2 and F3 generations were produced. The F3 generation included a polyploid plant with 2n = 32 that self-fertilized to yield fertile F4 offspring that could not mate with either of the parental species. These allopolyploid plants had the

same appearance and chromosome number as the naturally occurring G. tetrahit. When the experimentally produced plants were crossed with the naturally occurring G. tetrahit, a fertile F1 generation was formed. Thus, the experiment duplicated the speciation process that occurred in nature. Changing ecology causes sympatric speciation in animals Biologists have observed the occurrence of sympatric speciation in animals, but its significance—how often it happens and under what conditions—is still debated. Many examples of sympatric speciation in animals involve parasitic insects and rely on genetic mechanisms other than polyploidy. For example, in the 1860s in the Hudson River Valley of New York, a population of fruit maggot flies (Rhagoletis pomonella) parasitic on the small red fruits of native hawthorn trees was documented to have switched to a new host, domestic apples, which had been introduced from Europe. Although the sister populations (hawthorn maggot flies and apple maggot flies) continue to occupy the same geographic area, no gene flow occurs between them because they eat, mate, and lay their eggs on different hosts (FIG. 20-11). In other words, because the hawthorn and apple maggot flies have diverged and are reproductively isolated from each other, they have effectively become separate species. Most entomologists, however, still recognize hawthorn and apple maggot flies as a single species because their appearance is virtually identical.

Hawthorn maggot fly range

Primula floribunda

Primula kewensis

Primula verticillata

Apple maggot fly and hawthorn maggot fly range

FIGURE 20-11 Ranges of apple and hawthorn maggot flies FIGURE 20-10 Animated Sympatric speciation of a primrose A new species of primrose, Primula kewensis, arose in 1898 as an allopolyploid derived from the interspecific hybridization of P. floribunda and P. verticillata. Today, P. kewensis is a popular houseplant.

Apple and hawthorn maggot flies are sympatric throughout the northern half of the hawthorn maggot fly’s range. (Adapted from G. L. Bush, “Sympatric Host Race Formation and Speciation in Frugivorous Flies of the Genus Rhagoletis [Diptera, Tephritidae],” Evolution, Vol. 23, No. 2, June 1969.)

In situations such as that of the fruit maggot flies, a mutation arises in an individual and spreads through a small group of insects by sexual reproduction. The particular mutation leads to disruptive selection (see Chapter 19), in which both the old and new phenotypes are favored. Selection is disruptive because the original population is still favored when it parasitizes the original host, and the mutants are favored by a new ecological opportunity—in this case, to parasitize a different host species. Additional mutations may occur that cause the sister populations to diverge even further.

Loss of Cichlid Diversity in Lake Victoria. In recent years, many of

the cichlid species living in Lake Victoria have become extinct. The Nile perch, a voracious predator that was deliberately introduced into the lake in 1960 to stimulate the local fishing economy and whose population exploded in the 1980s, has been blamed for the extinction of most of the cichlid species that have disappeared. Biologists from the University of Leiden in the Netherlands observed, however, that species were also disappearing in habitats in the lake where the Nile perch is known to have little or no impact. Ole Seehausen and his colleagues measured environmental features at 22 localities with such habitats and found strong correlations between bright and diverse male colors and clear, well-lit water. Each species has its own distinct male coloration, which is used by females of that species to choose mates. In clear water, members of each species do not interbreed with members of other species. As nearby forests have been cut, soil erosion has made the water of Lake Victoria more turbid (cloudy). Agricultural practices in the area have also contributed fertilizer and sediment pollution. Seehausen demonstrated that as the water became more turbid from pollution, light could not penetrate effectively. He hypothesized that as a result, females could not distinguish males of their own species from males of closely related species. As already discussed, cichlids have evolved during the past 12,000 years. Their relative youth as species means they may not

a–c: Ole Seehausen, University of Bern

Cichlids in East Africa intrigue evolutionary biologists Biologists have studied the speciation of colorful fishes known as cichlids (pronounced “sik-lids”) in several East African lakes. The different species of cichlids in a given lake have remarkably different eating habits, which are reflected in the shapes of their jaws and teeth. Some graze on algae; some consume dead organic material at the bottom of the lake; and others are predatory and eat plankton (microscopic aquatic organisms), insect larvae, the scales off fish, or even other cichlid species. In some cichlids, food preferences are related to size (smaller cichlids consume plankton), which in turn is related to mating preference (small, plankton-eating cichlids mate only with other small, plankton-eating cichlids). In other cichlids, related species do not differ in size but only in color and mating habits. Each species has a distinct male coloration, which matches the preferences of females of that species in choosing mates (FIG. 20-12). Charles Darwin called choosing a mate based on its color or some other characteristic sexual selection. He recognized that female preference for showy colors or elaborate ornaments on males, if it was based on inherited variation, could result in these male sexual traits becoming more common and more pronounced over time. Female preference for a particular male trait confers a selective advantage to the males exhibiting that trait because they have an improved chance to mate and pass on their genes. Darwin suggested that sexual selection, like natural selection, could eventually result in the evolution of new species. (Sexual selection is discussed in greater detail in Chapter 52.) DNA sequence data indicate that the cichlid species within a single lake are more closely related to one another than to fishes in nearby lakes or rivers. These molecular data suggest that cichlid

species evolved sympatrically, or at least within the confines of a lake, rather than by repeated colonizations by fish populations in nearby rivers. How rapidly did sympatric speciation occur in cichlids? Seismic and drill core data provide evidence that Lake Victoria dried up completely during the Late Pleistocene, when much of north and equatorial Africa was arid. The diverse cichlid species may have arisen after the climate became wetter and Lake Victoria refilled. Further, the more than 500 endemic (found nowhere else) cichlid species in Lake Victoria may have evolved from one or a few common ancestors in a remarkably short period—as little as 12,000 years. Other biologists have offered competing hypotheses about the timing and geographic origin of Lake Victoria’s cichlids. Regardless of the uncertainties of certain details, Lake Victoria’s cichlids are widely recognized as exemplifying the fastest rate of evolution known for such a large group of vertebrate species.

(a) Pundamilia pundamilia males have bluish-silver bodies.

(b) Pundamilia nyererei males have red backs.

FIGURE 20-12 Color variation in Lake Victoria cichlids Some evidence suggests that changes in male coloration may be the first step in speciation of Lake Victoria cichlids. Later, other traits, including ecological characteristics, diverge. Female cichlids generally have cryptic coloration; their drab colors help them blend into their surroundings.

(c) Pundamilia “red head“ males have a red ”chest.“ (The ”red head“ species has not yet been scientifically named.)

yet have evolved additional reproductive isolating mechanisms other than mate preference based on color. It also means that closely related species can be expected to interbreed without loss of fertility. With increasing turbidity, males lose their bright colors because they are no longer favored by sexual selection, and females have mated with males of other species (FIGURE 20-13). Thus, many species are being replaced with a few hybrids.

The study of hybrid zones has made important contributions to what is known about speciation When two populations have significantly diverged as a result of geographic separation, there is no easy way to determine if the speciation process is complete (recall the disagreement about whether Porto Santo rabbits are a separate species or a subspecies). If such populations, subspecies, or species come into secondary contact before strong prezygotic isolating mechanisms have evolved, they may hybridize, producing fertile offspring, where they meet. A hybrid zone is an area of overlap between two recently diverged populations in which interbreeding takes place and hybrid offspring are common. Hybrid zones are typically narrow, presumably because the hybrids are not well adapted for either parental environment. The hybrid population is typically very small compared with the parental populations. On the Great Plains of North America, red-shafted and yellow-shafted flickers (types of woodpeckers) meet and interbreed. The red-shafted flicker, named for the male’s red underwings and tail, is found in the western part of North America, from the Great Plains to the Pacific Ocean. Yellow-shafted flicker males, which have yellow underwings and tails, range east of the Rock-

ies. Hybrid flickers, which form a stable hybrid zone from Texas to southern Alaska, are varied in appearance, although many have orange underwings and tails. According to the biological species concept, if red-shafted and yellow-shafted flickers are two species, they should maintain their reproductive isolation. Because a hybrid zone has formed, most biologists think the red-shafted and yellow-shafted flickers are geographic subspecies instead of separate species; red-shafted, yellow-shafted, and hybrid flickers are collectively called northern flickers. Three evolutionary processes may occur in hybrid zones When two diverging populations or closely related species come into contact and form a hybrid zone, one of three possibilities can occur: reinforcement, which helps maintain two separate species; fusion of the two species into one; or stability of the hybrid zone (FIG. 20-14). Reinforcement occurs when the hybrids in the hybrid zone are less fit than the two parental populations. Reinforcement is the increase in reproductive isolation that often occurs over time in a hybrid zone—that is, natural selection strengthens and increases the number of prezygotic barriers between the two species. As a result, fewer and fewer hybrids are formed, and the hybrid zone gradually disappears. In fusion, the hybrids in a hybrid zone are as fit as either of the parental populations. Over time, differences in the reproductive barriers between the two parental populations weaken, and the hybrid zone enlarges. The eventual result is a fusing of two species (or subspecies) into one. Hybridization of the Lake Victoria cichlids (discussed earlier in the chapter) is an example of species loss due to fusion.

KEY EXPERIMENT QUESTION: Why is cichlid diversity declining in Lake Victoria? HYPOTHESIS: Cichlids are disappearing from Lake Victoria because of changes in their habitat. EXPERIMENT: Seehausen and colleagues took field measurements in 22 different locations in Lake Victoria and later verified these field observations with laboratory experiments in which two cichlid species were placed in aquaria that were either well-lit or not.

RESULTS AND CONCLUSION: When aquaria were well-lit (left), females of two closely related cichlid species (one red, the other blue) consistently chose mates of their own species (circled area) over those of other species. When light was blocked to simulate turbid conditions found in parts of Lake Victoria (right), females often chose males of other species. Thus, increasing turbidity of Lake Victoria’s water is preventing females of cichlid species from using color to select mates of their own species. As a result, females are interbreeding with males of other species, and the number of cichlid species is declining as the distinctive gene pools of different species are merging. Source: Seehausen, O., et al. "Speciation through Sensory Drive in Cichlid Fish." Nature, Vol. 455, October 2, 2008.

FIGURE 20-13 Explaining the rapid loss of cichlid diversity in Lake Victoria

(a) Reinforcement. Hybrids are not as fit as the parents. Over time, the differences between the two parental populations increase, and hybrids are no longer produced. As the hybrid zone shrinks, the resulting outcome is two species.

ronments. In this case, the two incipient species are adapted to two different communities, and the hybrids’ combination of characters from both parental populations allows them to flourish in the ecotone. Eventually, a third species may form in such a hybrid zone. (Ecotones are discussed further in Chapter 56.) In the example of the flicker hybrid zone, reinforcement does not appear to be occurring. At the same time, the two kinds of flickers are not fusing together into one distinct group. The flicker hybrid zone has not expanded; the two types of flickers have maintained their distinctiveness and have not rejoined into a single, freely interbreeding population. Thus, at this time, flicker hybrids have formed a stable hybrid zone.

Review ■

■

■

What are five geographic barriers that might lead to allopatric speciation? How do hybridization and polyploidy cause a new plant species to form in as little as one generation? What is the likely mechanism of speciation for pupfishes? for cichlids?

20.4 THE RATE OF EVOLUTIONARY CHANGE ■ ■ LEARNING OBJECTIVE 5 (b) Fusion. Hybrids are as fit as the two parental populations. Over time, differences between the two parental populations weaken and the hybrid zone enlarges. The essential result is a single species.

(c) Stability. Hybrids are more fit than either parental population. The fitness of the hybrids results in a stable hybrid zone. Eventually, a third species may form in the hybrid zone.

FIGURE 20-14 How hybrid zones between two closely related species or diverging populations can change over time

Stability is an evolutionary process in which the hybrids are more fit than either parental population. Hybrids continue to be formed, resulting in a stable hybrid zone. Stable hybrid zones often occur in ecotones, areas of transition between two different envi-

Discuss the pace of evolution by describing punctuated equilibrium and phyletic gradualism.

We have seen that speciation is hard for us to directly observe as it occurs. Does the fossil record provide clues about how rapidly new species arise? Biologists have long recognized that the fossil record lacks many transitional forms; the starting points (ancestral species) and the end points (new species) are present, but the intermediate stages in the evolution from one species to another are often lacking. This observation has traditionally been explained by the incompleteness of the fossil record, although scientists have sometimes made new fossil discoveries that fill in the missing parts. Two different models—punctuated equilibrium and phyletic gradualism—explain evolution as observed in the fossil record (FIG. 20-15). The punctuated equilibrium model was proposed by paleontologists who questioned whether the fossil record really is as incomplete as it initially appeared. In the history of a species, long periods of stasis (little or no evolutionary change) appeared to be punctuated, or interrupted, by short periods of rapid speciation that were perhaps triggered by changes in the environment—that is, periods of great evolutionary stress. Thus, speciation proceeded in “spurts.” These relatively short periods of active evolution (perhaps 100,000 years) were followed by long periods (perhaps 2 million years) of stability. Punctuated equilibrium accounts for the abrupt appearance of a new species in the fossil record, with little or no evidence of intermediate forms. Proponents hypothesize that few transitional forms appear in the fossil record because few transitional forms occurred during speciation.

KEY POINT

Punctuated equilibrium and phyletic gradualism represent opposite ends of a spectrum of speciation patterns. Speciation often proceeds in an intermediate or mixed pattern.

Stasis

Extinction of original species

Slow, gradual changes Stasis

Stasis

Time

Time

Divergence is sudden, with rapid changes

Divergence is gradual

Stasis (little change)

Structural changes

(a) In punctuated equilibrium, long periods of stasis are interrupted by short periods of rapid speciation.

Structural changes

(b) In phyletic gradualism, a slow, steady change in species occurs over time.

FIGURE 20-15 Punctuated equilibrium and gradualism In this figure, structural changes in the lizards are represented by changes in skin color.

In contrast, the traditional Darwinian view of evolution espouses the phyletic gradualism model, in which evolution proceeded continuously over long periods. Gradualism is rarely observed in the fossil record because the record is incomplete. (Recall from Chapter 18 that precise conditions are required for fossil formation. Most organisms leave no trace of their existence because they decompose when they die.) Occasionally, a complete fossil record of transitional forms has been discovered and cited as a strong case for gradualism. The gradualism model maintains that populations slowly diverge from one another by the gradual accumulation of adaptive characteristics within each population. These adaptive characteristics accumulate as a result of different selective pressures encountered in different environments. Scientists supporting phyletic gradualism maintain that any periods of stasis evident in the fossil record are the result of stabilizing selection (see Chapter 19). They also emphasize that stasis in fossils is deceptive because fossils do not reveal all aspects of evolution. Although fossils display changes in external structure and skeletal structure, other genetic changes—in physiology, internal structure, resistance to infection, and behavior—all of which also represent evolution, are not evident. Gradualists recognize rapid evolution only when strong directional selection occurs. Many biologists today embrace both models to explain the fossil record. They contend that the pace of evolution may be abrupt in certain instances and gradual in others and that neither punctuated

equilibrium nor gradualism exclusively characterizes the changes associated with evolution. Many biologists do not view the distinction between punctuated equilibrium and phyletic gradualism as real. They suggest that genetic changes occur gradually and at a roughly constant pace and that the majority of these mutations do not cause speciation. When the mutations that do cause speciation occur, they are dramatic and produce a pattern consistent with the punctuated equilibrium model.

Review ■

Are the phyletic gradualism and punctuated equilibrium models mutually exclusive? Explain your answer.

20.5 MACROEVOLUTION ■ ■ LEARNING OBJECTIVES 6 Define macroevolution. 7 Discuss macroevolution in the context of novel features, including pre8

adaptations, allometric growth, and paedomorphosis. Discuss the macroevolutionary significance of adaptive radiation and extinction.

Macroevolution is large-scale phenotypic changes in populations that generally warrant their placement in taxonomic groups at the

Evolutionary novelties originate through modifications of pre-existing structures New designs arise from structures already in existence. A change in the basic pattern of an organism produces something unique, such as wings on insects, flowers on plants, and feathered wings on birds. Usually, these evolutionary novelties are variations of some pre-existing structures, called preadaptations, that originally fulfilled one role but subsequently changed in a way that was adaptive for a different role. Feathers, which evolved from reptilian scales and may have originally provided thermal insulation in primitive birds and some dinosaurs, represent a preadaptation for flight. With gradual modification, feathers evolved to function in flight as well as to fulfill their original thermoregulatory role. (Interestingly, a few feather-footed bird species exist; this phenotype is the result of a change in gene regulation that alters scales, normally found on bird feet, into feathers.) How do such evolutionary novelties originate? Many are probably due to changes during development. Regulatory genes exert control over hundreds of other genes during development, and very slight genetic changes in regulatory genes could ultimately cause major structural changes in the organism (see Chapters 17 and 18). For example, during development most organisms exhibit varied rates of growth for different parts of the body, known as allometric growth (from the Greek allo, “different,” and metr, “measure”). The size of the head in human newborns is large in proportion to the rest of the body. As a human grows and matures, the torso, hands, and legs grow more rapidly than the head. Allometric growth is found in many organisms, including the male fiddler crab with its single, oversized claw, and the ocean sunfish with its enlarged tail (FIG. 20-16). If growth rates are altered even slightly, drastic changes in the shape of an organism may result, changes that may or may not be adaptive. For example, allometric growth may help explain the extremely small and relatively useless forelegs of the dinosaur Tyrannosaurus rex, compared with those of its ancestors. Sometimes novel evolutionary changes occur when a species undergoes changes in the timing of development. Consider, for example, the changes that would occur if juvenile characteristics were retained in the adult stage, a phenomenon known as paedomorphosis (from the Greek paed, “child,” and morph, “form”). Adults of some salamander species have external gills and tail fins, features found only in the larval (immature) stages of other salamanders.

Tail

approx. 1 mm Newly hatched ocean sunfish

(a) A newly hatched ocean sunfish, only 1 mm long, has an extremely small tail.

Adult ocean sunfish

(b) This allometric transformation is visualized by drawing rectangular coordinate lines through a picture of the juvenile fish and then changing the coordinate lines mathematically.

Richard Herrmann

species level and higher—that is, new species, genera, families, orders, classes, and even phyla, kingdoms, and domains. One concern of macroevolution is to explain evolutionary novelties, which are large phenotypic changes such as the appearance of jointed limbs during the evolution of arthropods (crustaceans, insects, and spiders). These phenotypic changes are so great that the new species possessing them are assigned to different genera or higher taxonomic categories. Studies of macroevolution also seek to discover and explain major changes in species diversity through time such as occur during adaptive radiation, when many species appear, and mass extinction, when many species disappear. Thus, evolutionary novelties, adaptive radiation, and mass extinction are important aspects of macroevolution.

(c) An ocean sunfish swims off the coast of southern California. The adult ocean sunfish may reach 4 m (13 ft) and weigh about 1500 kg (3300 lb).

FIGURE 20-16 Allometric growth in the ocean sunfish The tail end of an ocean sunfish (Mola mola) grows faster than the head end, resulting in the unique shape of the adult.

Retention of external gills and tail fins throughout life obviously alters the salamander’s behavioral and ecological characteristics (FIG. 20-17). Perhaps such salamanders succeeded because they had a selective advantage over “normal” adult salamanders; that is, by remaining aquatic, they did not have to compete for food with the terrestrial adult forms of related species. The paedomorphic forms also escaped the typical predators of terrestrial salamanders (although they had other predators in their aquatic environment). Studies suggest that paedomorphosis in salamanders is probably the result of mutations in genes that block the production of hormones that stimulate metamorphic changes. When paedomorphic salamanders receive hormone injections, they develop into adults lacking external gills and tail fins.

Stephen Dalton/Photo Researchers, Inc.

FIGURE 20-17 Paedomorphosis in a salamander An adult axolotl salamander (Ambystoma mexicanum) retains the juvenile characteristics of external gills (feathery structures protruding from the neck) and a tail fin (not visible). Paedomorphosis allows the axolotl to remain permanently aquatic and to reproduce without developing typical adult characteristics.

Adaptive radiation is the diversification of an ancestral species into many species Some ancestral species have given rise to far more species than have other evolutionary lineages. Adaptive radiation is the evolutionary diversification of many related species from one or a few ancestral species in a relatively short period. Evolutionary innovations such as those we considered in the last section may trigger an adaptive radiation. Alternatively, an adaptive radiation may occur because the ancestral species happens to have been in the right place at the right time to exploit numerous ecological opportunities. Biologists developed the concept of adaptive zones to help explain this type of adaptive radiation. Adaptive zones are new ecological opportunities that were not exploited by an ancestral organism. At the species level, an adaptive zone is essentially identical to one or more similar ecological niches (the functional roles of species within a community; see Chapter 54). Examples of adaptive zones include nocturnal flying to catch small insects, grazing on grass while migrating across a savanna, and swimming at the ocean’s surface to filter out plankton. When many adaptive zones are empty, as they were in Lake Victoria when it refilled some 12,000 years ago (discussed earlier in the chapter), colonizing species such as the cichlids may rapidly diversify and exploit them. Because islands have fewer species than do mainland areas of similar size, latitude, and topography, vacant adaptive zones are more common on islands than on continents. Consider the Hawaiian honeycreepers, a group of related birds found in the Hawaiian Islands. When the honeycreeper ancestors reached Hawaii, few other birds were present. The succeeding generations of honeycreepers quickly diversified into many new species and, in the process, occupied the many available adaptive zones that on the mainland are occupied by finches, honeyeaters, treecreepers, and woodpeckers. The diversity of their bills is a particularly good illus-

tration of adaptive radiation (FIG. 20-18). Some honeycreeper bills are curved to extract nectar out of tubular flowers, whereas others are short and thick for ripping away bark in search of insects. Another example of vacant adaptive zones involves the Hawaiian silverswords, 28 species of closely related plants found only in the Hawaiian Islands. When the silversword ancestor, a California plant related to daisies, reached the Hawaiian Islands, many diverse environments, such as cool, arid mountains; exposed lava flows; dry woodlands; shady, moist forests; and wet bogs were present and more or less unoccupied. The succeeding generations of silverswords quickly diversified in structure and physiology to occupy the many adaptive zones available to them. The diversity in their leaves, which changed during the course of natural selection as different populations adapted to various levels of light and moisture, is a particularly good illustration of adaptive radiation (FIG. 20-19). Leaves of silverswords that are adapted to shady, moist forests are large, for example, whereas those of silverswords living in arid areas are small. The leaves of silverswords living on exposed volcanic slopes are covered with dense, silvery hairs that may reflect some of the intense ultraviolet radiation off the plant. Adaptive radiation appears more common during periods of major environmental change, but it is difficult to determine how these changes are related to adaptive radiation. It is possible that major environmental change indirectly affects adaptive radiation by increasing the extinction rate. Extinction produces empty adaptive zones, which provide new opportunities for species that remain. Mammals, for example, diversified and exploited a variety of adaptive zones relatively soon after the dinosaurs’ extinction. Flying bats, running gazelles, burrowing moles, and swimming whales all originated from early mammals that had coexisted with reptiles for millions of years.

Extinction is an important aspect of evolution Extinction, the end of a lineage, occurs when the last individual of a species dies. The loss is permanent, for once a species is extinct, it never reappears. Extinctions have occurred continually since the origin of life; by one estimate, only 1 species is alive today for every 2000 that have become extinct. Extinction is the eventual fate of all species, in the same way that death is the eventual fate of all individual organisms. Although extinction has a negative short-term impact on the number of species, it facilitates evolution over a period of thousands to millions of years. As mentioned previously, when species become extinct, their adaptive zones become vacant. Consequently, surviving species are presented with new evolutionary opportunities and may diverge, filling in some of the unoccupied zones. In other words, the extinct species may eventually be replaced by new species. During the long history of life, extinction appears to have occurred at two different rates. The continuous, low-level extinction of species is sometimes called background extinction. In contrast, five or possibly six times during Earth’s history, mass extinctions of numerous species and higher taxonomic groups have taken place in both terrestrial and marine environments. The most recent mass extinction, which occurred about 65 million years ago (mya), killed off many marine organisms, terrestrial plants, and

KEY POINT

Adaptive radiation occurs when a single ancestral species diversifies into a variety of species, each adapted to a different ecological niche.

Rips away bark to find insects Maui parrot bill

Kauai

Sips flower nectar ‘I‘iwi Oahu

Forages among leaves and branches Maui creeper Maui

Chisels holes in bark to get insects Akiapolaau Extinct Sipped flower nectar Black mamo

Extinct Habits unknow n Ula-ai-hawane

Hawaii

Picks food fr om cracks in the bark Akialoa

Sips flower nectar Apapane Feeds on snails and invertebrates Poo-uli

Sips flower nectar Crested honeycreeper

FIGURE 20-18 Adaptive radiation in Hawaiian honeycreepers Compare the various beak shapes and methods of obtaining food. Many honeycreeper species are now extinct or nearing extinction as a result of human activities, including the destruction of habitat and the introduction of predators such as rats, dogs, and pigs.

vertebrates, including the last of the dinosaurs. The time span over which a mass extinction occurs may be several million years, but that is relatively short compared with the 3.5 billion years or so of Earth’s history of life. Each period of mass extinction has been followed by a period of adaptive radiation of some of the surviving groups. The causes of past episodes of mass extinction are not well understood. Both environmental and biological factors seem to have been involved. Major changes in climate could have adversely affected those plants and animals that lacked the genetic flexibility to adapt. Marine organisms, in particular, are adapted to a steady, unchanging climate. If Earth’s temperature were to increase or decrease by just a few degrees overall, many marine species would probably perish. It is also possible that past mass extinctions were due to changes in the environment induced by catastrophes. If a large comet or small asteroid collided with Earth, the dust ejected into the atmosphere on impact could have blocked much of the sunlight. In addition to disrupting the food chain by killing many plants (and

therefore terrestrial animals), this event would have lowered Earth’s temperature, leading to the death of many marine organisms. Evidence exists that the extinction of dinosaurs was caused by an extraterrestrial object’s collision with Earth (see Chapter 21). Biological factors also trigger extinction. Competition among species may lead to the extinction of species that cannot compete effectively. The human species, in particular, has had a profound impact on the rate of extinction. The habitats of many animal and plant species have been altered or destroyed by humans, and habitat destruction can result in a species’s extinction. Some biologists think we have entered the greatest mass extinction episode in Earth’s history. (Extinction is discussed further in Chapter 57.)

Is microevolution related to speciation and macroevolution? The concepts presented in Chapters 18 and 19 represent the modern synthesis, in which mutation provides the genetic variation on which natural selection acts. The modern synthesis combines

(b) This silversword species (Wilkesia gymnoxiphium), which superficially resembles a yucca, is found only along the slopes of Waimea Canyon on the island of Kauai.

Jack Jeffrey Photography

Jack Jeffrey Photography

Jack Jeffrey Photography

(a) The Haleakala silversword (Argyroxyphium sandwicense ssp. macrocephalum) is found only in the cinders on the upper slope of Haleakala Crater on the island of Maui. This plant is adapted to low precipitation and high levels of ultraviolet radiation.

(c) Daubautia scabra is a small, herbaceous silversword found in moist to wet environments on several Hawaiian islands. (The fern fronds in the background give an idea of the small size of D. scabra.)

FIGURE 20-19 Adaptive radiation in Hawaiian silverswords The 28 silversword species, which are classified in three closely related genera, live in a variety of habitats.

Darwin’s theory with important aspects of genetics. Many aspects of the modern synthesis have been tested and verified at the population and subspecies levels. Many biologists contend that microevolutionary processes explain macroevolution. They assert that natural selection, mutation, genetic drift, and gene flow account for not only the genetic variation within populations but also the origin of new species and higher taxa as well. A considerable body of data from many fields supports the modern synthesis as it relates to speciation and macroevolution. Consider, for example, the fish–tetrapod transition (the evolution of four-legged amphibians from fishes), which was a major macroevolutionary event in the history of vertebrates (see Chapter 32). The fish–tetrapod transition is well documented in the fossil record. Studies of fossil intermediates show that the transition from aquatic fishes to terrestrial amphibians occurred as evolutionary novelties, such as changes in the limbs and skull roof, were added. These novelties accumulated as a succession of small changes over a period of 9 million to 14 million years. This time scale is sufficient to have allowed natural selection and other microevolutionary processes to produce the novel characters. Although few biologists doubt the role of natural selection and microevolution in generating specific adaptations, some question the extent of microevolution’s role in the overall pattern of life’s

history. These biologists ask whether speciation and macroevolution have been dominated by microevolutionary processes or by external, chance events, such as an impact by an asteroid. Chance events do not “care” about adaptive superiority but instead lead to the random extinction or survival of species. In the case of an asteroid impact, for example, certain species may survive because they were “lucky” enough to be in a protected environment at the time of impact. If chance events have been the overriding factor during life’s history, then microevolution cannot be the exclusive explanation for the biological diversity present today (see discussion of the role of chance in evolution in Chapter 18).

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Why are evolutionary novelties important to scientists studying macroevolution? What is a preadaptation? What is paedomorphosis? What roles do extinction and adaptive radiation play in macroevolution?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

20.1 (page 427) 1 Describe the biological species concept and list two potential problems with the concept. According to the biological species concept, a species consists of one or more populations whose members interbreed in nature to produce fertile offspring and do not interbreed with members of different species. ■ One problem with the biological species concept is that it applies only to sexually reproducing organisms. Also, although individuals assigned to different species do not normally interbreed, they may occasionally successfully interbreed. ■

20.2 (page 428) 2 Explain the significance of reproductive isolating mechanisms and distinguish among the various prezygotic and postzygotic barriers. Reproductive isolating mechanisms restrict gene flow between species. ■ Prezygotic barriers are reproductive isolating mechanisms that prevent fertilization from taking place. Temporal isolation occurs when two species reproduce at different times of the day, season, or year. In habitat isolation, two closely related species live and breed in different habitats in the same geographic area. In behavioral isolation, distinctive courtship behaviors prevent mating between species. Mechanical isolation is due to incompatible structural differences in the reproductive organs of similar species. In gametic isolation, gametes from different species are incompatible because of molecular and chemical differences. ■

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Postzygotic barriers are reproductive isolating mechanisms that prevent gene flow after fertilization has taken place. Hybrid inviability is the death of interspecific embryos during development. Hybrid sterility prevents interspecific hybrids that survive to adulthood from reproducing successfully. Hybrid breakdown prevents the offspring of hybrids that survive to adulthood and successfully reproduce from reproducing beyond one or a few generations. Explore reproductive isolation by clicking on the figures in CengageNOW.

20.3 (page 430) 3 Explain the mechanism of allopatric speciation and give an example. ■

Speciation is the evolution of a new species from an ancestral population. Allopatric speciation (for example, in Death Valley pupfishes) occurs when one population becomes geographically isolated from the rest of the species and subsequently diverges. Speciation is more likely to occur if the original isolated population is small because genetic drift is more significant in small populations. Explore allopatric speciation by clicking on the figures in CengageNOW.

4 Explain the mechanisms of sympatric speciation and give both plant and animal examples. Sympatric speciation does not require geographic isolation. In plants, sympatric speciation can result from allopolyploidy, in

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which a polyploid individual (one with more than two sets of chromosomes) is a hybrid derived from two species. One example of sympatric speciation by allopolyploidy is the hemp nettle. Sympatric speciation occurs in animals, such as fruit maggot flies and cichlids, but how often it occurs and under what conditions remain to be determined. Explore sympatric speciation by clicking on the figure in CengageNOW.

20.4 (page 438) 5 Discuss the pace of evolution by describing punctuated equilibrium and phyletic gradualism. ■ According to the punctuated equilibrium model, evolution of species proceeds in spurts. Short periods of active speciation are interspersed with long periods of stasis. According to the phyletic gradualism model, populations slowly diverge from one another by the accumulation of adaptive characteristics within a population.

20.5 (page 439) 6 Define macroevolution. Macroevolution is large-scale phenotypic changes in populations that typically warrant the placement of the populations in taxonomic groups at the species level and higher—that is, new species, genera, families, orders, classes, and even phyla, kingdoms, and domains. 7 Discuss macroevolution in the context of novel features, including preadaptations, allometric growth, and paedomorphosis. ■ Evolutionary novelties may be due to changes during development, the orderly sequence of events that occurs as an organism grows and matures. Slight genetic changes in regulatory genes could cause major structural changes in an organism. ■ Evolutionary novelties may originate from preadaptations, structures that originally fulfilled one role but changed in a way that was adaptive for a different role. Feathers are an example of a preadaptation. ■ Allometric growth, varied rates of growth for different parts of the body, results in overall changes in the shape of an organism. Examples include the expanded tail region of the ocean sunfish and the enlarged claw of the male fiddler crab. ■ Paedomorphosis, the retention of juvenile characteristics in the adult, occurs because of changes in the timing of development. Adult axolotl salamanders, with external gills and tail fins, are an example of paedomorphosis. 8 Discuss the macroevolutionary significance of adaptive radiation and extinction. ■ Adaptive radiation is the process of diversification of an ancestral species into many new species. Adaptive zones are new ecological opportunities that were not exploited by an ancestral organism. When many adaptive zones are empty, colonizing species may rapidly diversify and exploit them. Hawaiian honeycreepers and silverswords both underwent adaptive radiation after their ancestors colonized the Hawaiian Islands. ■

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Extinction is the death of a species. When species become extinct, the adaptive zones that they occupied become vacant, allowing other species to evolve and fill those zones. Background extinction

is the continuous, low-level extinction of species. Mass extinction is the extinction of numerous species and higher taxonomic groups in both terrestrial and marine environments.

T E S T YO U R U N D E R S TA N D I N G 1. A prezygotic barrier prevents (a) the union of egg and sperm (b) reproductive success by an interspecific hybrid (c) the development of the zygote into an embryo (d) allopolyploidy from occurring (e) changes in allometric growth 2. The reproductive isolating mechanism in which two closely related species live in the same geographic area but reproduce at different times is (a) temporal isolation (b) behavioral isolation (c) mechanical isolation (d) gametic isolation (e) hybrid inviability 3. Interspecific hybrids, if they survive, are (a) always sterile (b) always fertile (c) usually sterile (d) usually fertile (e) never sterile 4. The first step leading to allopatric speciation is (a) hybrid inviability (b) hybrid breakdown (c) adaptive radiation (d) geographic isolation (e) paedomorphosis 5. The pupfishes in the Death Valley region are an example of which evolutionary process? (a) background extinction (b) allopatric speciation (c) sympatric speciation (d) allopolyploidy (e) paedomorphosis 6. Which of the following evolutionary processes is associated with allopolyploidy? (a) gradualism (b) allometric growth (c) sympatric speciation (d) mass extinction (e) preadaptation

7. According to the punctuated equilibrium model, (a) populations slowly diverge from one another (b) the evolution of species occurs in spurts interspersed with long periods of stasis (c) evolutionary novelties originate from preadaptations (d) reproductive isolating mechanisms restrict gene flow between species (e) the fossil record, being incomplete, does not accurately reflect evolution as it actually occurred 8. The evolutionary conversion of reptilian scales into feathers is an example of (a) allometric growth (b) paedomorphosis (c) gradualism (d) hybrid breakdown (e) a preadaptation 9. Adaptive radiation is common following a period of mass extinction, probably because (a) the survivors of a mass extinction are remarkably well adapted to their environment (b) the unchanging environment following a mass extinction drives the evolutionary process (c) many adaptive zones are empty (d) many ecological niches are filled (e) the environment induces changes in the timing of development for many species 10. The Hawaiian silverswords are an excellent example of which evolutionary process? (a) allometry (b) preadaptation (c) microevolution (d) adaptive radiation (e) extinction

CRITICAL THINKING 1. EVOLUTION LINK. Why is allopatric speciation more likely to occur if the original isolated population is small?

Harry Engels/Photo Researchers, Inc.

2. EVOLUTION LINK. The photograph shows a “zebrass,” a sterile hybrid produced on a Texas ranch by a cross between a zebra and a donkey. Based on what you have learned about prezygotic and postzygotic isolating mechanisms, which reproductive isolating mechanism(s) would you say is/are probably at work here? How would you test your hypothesis?

3. EVOLUTION LINK. Could hawthorn and apple maggot flies be considered an example of assortative mating, which was discussed in Chapter 19? Explain your answer. 4. EVOLUTION LINK. Because mass extinction is a natural process that may facilitate evolution during the period of thousands to millions of years that follow it, should humans be concerned about the current mass extinction that we are causing? Why or why not? 5. ANALYZING DATA. Examine Figure 20-2c, and predict which date is likeliest for researchers to have collected wood and leopard frogs and interbred them in the lab: March 10, April 1, or April 15. Explain your answer. Additional questions are available in CengageNOW at www.cengage.com/ login.

21

The Origin and Evolutionary History of Life

© Phil Degginger/Carnegie Museum/Alamy

Fossil of an Early Permian reptile. This well-preserved fossil of Orobates pabsti was matched to well-preserved fossil footprints found nearby. Although the limbs are spread out in death, Orobates walked with an upright gait with its limbs under its body. Discovered and photographed in Germany.

KEY CONCEPTS 21.1 Although there is no direct fossil evidence of the origin of life, biochemical experiments have demonstrated how the complex organic molecules that are found in all living organisms may have formed. 21.2 The first cells were probably heterotrophic, anaerobic, and prokaryotic. Photosynthesis, aerobic respiration, and eukaryotic cell structure represent several major advances that occurred during the early history of life.

T

he preceding three chapters were concerned with biological evolution. However, we have not dealt with what many regard as a funda-

mental question: How did life begin? Although biologists generally accept the hypothesis that life developed from nonliving matter, exactly how this process, called chemical evolution, occurred is not certain. Current models suggest that small organic molecules first formed spontaneously and accumulated over time. Large organic macromolecules such as proteins and nucleic acids could have then assembled from the smaller molecules. The macromolecules interacted, combining into more complicated

21.3 The fossil record tells us much of what we know

structures that could eventually metabolize and replicate, passing their

about the history of life, such as what kinds of organisms existed and where and when they lived. Scientists identify and demonstrate relationships among fossils in rock layers from different periods of geologic time.

genetic information on to their descendants, which eventually became the first true cells. After the first cells originated, they diverged over several billion years into the rich biological diversity that characterizes our planet today. Initially, unicellular prokaryotes (archaea and bacteria) predominated, followed by unicellular eukaryotes. The first multicellular eukaryotes were soft-bodied marine animals that did not leave many fossils.

Shelled animals and other marine invertebrates (animals without backbones) appeared next, as exemplified by trilobites, primitive aquatic arthropods. Marine invertebrates were followed by the first vertebrates (animals with backbones). The first fishes with jaws appeared and diversified. Some of these gave rise to amphibians, the first vertebrates with limbs that were capable of moving about on land. Amphibians gave rise to reptiles, which diversified and populated the land (see photograph). Reptiles, in turn, gave rise independently to birds and to mammals. Plants underwent a comparable evolutionary history and diversification. In this chapter we survey life over a vast span of time, starting when our planet was relatively young. We first examine proposed models about how life began and then trace life’s long evolutionary history.

21.1 CHEMICAL EVOLUTION ON EARLY EARTH ■ ■ LEARNING OBJECTIVES 1 2

Describe the conditions that scientists think existed on early Earth. Contrast the prebiotic soup hypothesis and the iron–sulfur world hypothesis.

Scientists generally agree that life’s beginnings occurred under environmental conditions quite different from those of today. Therefore,

Raw materials from meteorites

we must examine the conditions of early Earth to understand the origin of life. Evidence from many sources provides us with clues that help us formulate plausible scenarios of the steps in which life originated. Study of the origin of life is an active area of scientific research today, and many important contributions are adding to our understanding of how life began. Astrophysicists and geologists estimate that Earth is about 4.6 billion years old. The atmosphere of early Earth apparently included carbon dioxide (CO2), water vapor (H2O), carbon monoxide (CO), and nitrogen (N2). It may also have contained some ammonia (NH3), hydrogen sulfide (H2S), and methane (CH4), although ultraviolet radiation from the sun most likely broke these reduced molecules down rapidly. The early atmosphere probably contained little or no free oxygen (O2). Four requirements must have existed for the chemical evolution of life: little or no free oxygen, a source of energy, the availability of chemical building blocks, and time. First, life could have begun only in the absence of free oxygen. Oxygen is quite reactive and would have oxidized the organic molecules that are necessary building blocks in the origin of life. Earth’s early atmosphere was probably strongly reducing, which means that any free oxygen would have reacted with other elements to form oxides. Thus, oxygen would have been tied up in compounds. The origin of life would also have required energy to do the work of building biological molecules from simple inorganic chemicals. Early Earth was a place of high energy with violent thunderstorms; widespread volcanic activity; bombardment from meteorites and other extraterrestrial objects; and intense radiation, including ultraviolet radiation from the sun (FIG. 21-1). The young

Raw materials from volcanoes

Energy from lightning

Cosmic and ultraviolet radiation

Land Ocean

Raw materials from deep-sea thermal vents

FIGURE 21-1 Conditions on early Earth Volcanoes erupted, spewing gases that contributed to the atmosphere, and violent thunderstorms triggered torrential rainfall that washed molecules out of the atmosphere and eroded the land. Meteorites and other extraterrestrial objects bombarded Earth, cataclysmically changing the crust, ocean, and atmosphere. Deepsea thermal vents, heated by Earth’s internal heat, spewed forth chemicals. High-energy radiation from cosmic and ultraviolet rays blasted the planet.

sun probably produced more ultraviolet radiation than it does today, and ancient Earth had no protective ozone layer to filter it. A third requirement would have been the presence of the chemical building blocks needed for chemical evolution. These included water, dissolved inorganic minerals (present as ions), and the gases present in the early atmosphere. A final requirement for the origin of life was time for molecules to accumulate and react with one another. Earth is approximately 4.6 billion years old, and evidence suggests that life arose early in Earth’s history.

KEY EXPERIMENT QUESTION: Could organic molecules have formed in the conditions of early Earth? HYPOTHESIS: Organic molecules can form in a reducing atmosphere similar to that thought to be present on early Earth. EXPERIMENT: The apparatus that Miller and Urey used to simulate the reducing atmosphere of early Earth contained hydrogen, methane, ammonia, and water. An electrical spark was produced in the upper right flask to simulate lightning. Electrodes

Organic molecules formed on primitive Earth Because organic molecules are the building materials for organisms, it is reasonable to first consider how they may have originated. Two main models seek to explain how the organic precursors of life originated: the prebiotic soup hypothesis proposes that these molecules formed near Earth’s surface, whereas the iron– sulfur world hypothesis proposes that organic precursors formed at cracks in the ocean’s floor.

NH3 CH4 To vacuum

H2O

H2 Spark chamber

Organic molecules may have been produced at Earth’s surface

The concept that simple organic molecules such as sugars, nucleotide bases, and amino acids could form spontaneously from simpler raw materials was first advanced in the 1920s by two scientists working independently: A. I. Oparin, a Russian biochemist, and J. B. S. Haldane, a Scottish physiologist and geneticist. Their hypothesis was tested in the 1950s by U.S. biochemists Stanley Miller and Harold Urey, who designed a closed apparatus that simulated conditions that presumably existed on early Earth (FIG. 21-2). They exposed an atmosphere rich in H2, CH4, H2O, and NH3 to an electric discharge that simulated lightning. Their analysis of the chemicals produced during one week revealed that amino acids and other organic molecules had formed. Although more recent data suggest that Earth’s early atmosphere was not rich in methane, ammonia, or hydrogen, similar experiments using different combinations of gases have produced a wide variety of organic molecules that are important in contemporary organisms. These include all 20 amino acids, several sugars, lipids, the nucleotide bases of RNA and DNA, and ATP (when phosphate is present). Thus, before life began, its chemical building blocks were probably accumulating as a necessary step in chemical evolution. Oparin envisioned that the organic molecules would, over vast spans of time, accumulate in the shallow seas to form a “sea of organic soup.” He envisioned smaller organic molecules (monomers) combining to form larger ones (polymers) under such conditions. Evidence gathered since Oparin’s time indicates that organic polymers may have formed and accumulated on rock or clay surfaces rather than in the primordial seas. Clay, which consists of microscopic particles of weathered rock, is particularly intriguing as a possible site for early polymerizations because it binds organic monomers and contains zinc and iron ions that may have served as catalysts. Laboratory experiments have confirmed that organic polymers form spontaneously from monomers on hot rock or clay surfaces.

Condenser

Input gases

Boiling chamber for gases and water Heat source

Organic molecules collect in the trap

RESULTS AND CONCLUSION: The gases present in the flask reacted together, and in one week a variety of simple organic compounds, such as amino acids, accumulated in the trap at the bottom. Thus, the formation of organic molecules—the first step in the origin of life—can be produced from simple precursors. Source: Miller, S. L., and H. C. Urey. “Organic Compound Synthesis on the Primitive Earth.” Science, Vol. 130, July 1959.

FIGURE 21-2 Animated Miller and Urey’s experiment

in chemical evolution

Organic molecules may have formed at hydrothermal vents Some biologists hypothesize that early polymerizations leading to the origin of life may have occurred in cracks in the deep-ocean floor where hot water, carbon monoxide, and minerals such as sulfides of iron and nickel spew forth—the iron–sulfur world hypoth-

Steven Brooke and Richard LeDuc

esis. Such hydrothermal vents would have been better protected than Earth’s surface from the catastrophic effects of meteorite bombardment. Today, these anaerobic hot springs produce precursors of biological molecules and of energy-rich “food,” including the highly reduced compounds hydrogen sulfide and methane. These chemicals support a diverse community of microorganisms, clams, crabs, tube worms, and other animals (see Inquiring About: Life without the Sun, in Chapter 55). Testing the iron–sulfur world hypothesis at hydrothermal vents is difficult, but laboratory experiments simulating the high pressures and temperatures at the vents have yielded intriguing results. For example, iron and nickel sulfides catalyze reactions between carbon monoxide and hydrogen sulfide, producing acetic acid and other simple organic compounds. Also, experiments show that ammonia, one of the precursors of proteins and nucleic acids, is produced in abundance, suggesting that vents were ammonia-rich environments in the prebiotic world and that the earliest organisms could have formed in this seemingly inhospitable environment.

2 μm

FIGURE 21-3 Microspheres These tiny protobionts exhibit some of the properties of life.

Review ■

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What are the four requirements for chemical evolution, and why is each essential? What is the prebiotic soup hypothesis? How does the iron–sulfur world hypothesis differ from the prebiotic soup hypothesis?

21.2 THE FIRST CELLS ■ ■ LEARNING OBJECTIVES 3 4 5

Outline the major steps hypothesized to have occurred in the origin of cells. Explain how the evolution of photosynthetic autotrophs affected both the atmosphere and other organisms. Describe the hypothesis of serial endosymbiosis.

After the first polymers formed, could they have assembled spontaneously into more complex structures with an outer membranous boundary? Scientists have synthesized several different protobionts, which are vesicle-like assemblages of abiotically produced (that is, not produced by organisms) organic polymers. These protobionts resemble living cells in several ways, thus providing clues as to how aggregations of complex nonliving molecules took that “giant leap” and became living cells. These protobionts exhibit many functional and structural attributes of living cells. They often divide in half (binary fission) after they have sufficiently “grown.” Protobionts maintain an internal chemical environment that is different from the external environment (homeostasis), and some of them show the beginnings of metabolism (catalytic activity). They are highly organized, considering their relatively simple composition. Microspheres are a type of protobiont formed by adding water to abiotically formed polypeptides (FIG. 21-3). Some microspheres are excitable: they produce an electrical potential across their surfaces, reminiscent of electrochemical gradients in cells. Microspheres can also absorb materials from their surroundings

(selective permeability) and respond to changes in osmotic pressure as though membranes enveloped them, even though they contain no lipid. In 2008, scientists constructed protobionts in which membranes of fatty acids and monoglycerides surround larger DNA-like molecules. These protobionts are impermeable to the genetic molecules but allow smaller organic molecules—that is, “nutrients”— to enter. This study is significant because it suggests how early cells could have held on to their “genes” and absorbed needed nutrients from their environment without the sophisticated membrane structure associated with contemporary cells. The study of protobionts shows that relatively simple nonliving structures have some of the properties of life. However, it is a major step (or several steps) to go from simple molecular aggregates such as protobionts to living cells.

The origin of a simple metabolism within a membrane boundary may have occurred early in the evolution of cells Although we have learned many things about how organic molecules may have formed on primitive Earth, the problem of how pre-cells evolved into living cells remains to be solved. Let’s examine one possible scenario—the metabolism first hypothesis— supported by chemist Robert Shapiro at New York University and many others. Shapiro envisions life beginning as a self-sustaining, organized system consisting of chemical reactions between simple molecules that are enclosed within a boundary—that is, separated from the external environment (FIG. 21-4a). A source of energy must be available to drive these chemical reactions (thereby maintaining an internal organization different from the external environment). To be self-sustaining, the energy source must be coupled to the chemical reaction sequence (FIG. 21-4B). (Recall how ATP links exergonic and endergonic reactions, shown in Figure 7-6.)

As the pre-cells continue to evolve, their organization increases (FIG. 21-4c). Finally, to survive, the pre-cell system must grow and reproduce. One of the most significant parts of the origin of living cells from pre-cells was the evolution of molecular reproduction (FIG. 21-4d).

Molecular reproduction was a crucial step in the origin of cells (a) Within a membranous boundary, chemical reactions occur among simple molecules.

(b) A source of energy is coupled to the chemical reaction sequence.

In living cells, genetic information is stored in the nucleic acid DNA, which is transcribed into messenger RNA (mRNA), which in turn is translated into the proper amino acid sequence in proteins. All three macromolecules in the DNA ¡ RNA ¡ protein sequence contain precise information. Of these, only DNA and RNA are capable of self-replication (although only in the presence of the proper enzymes in cells today). Because both RNA and DNA can form spontaneously on clay in much the same way that other organic polymers do, the question becomes, which molecule, DNA or RNA, first appeared in the prebiotic world? Many scientists have suggested that RNA was the first informational molecule to evolve in the progression toward a selfsustaining, self-reproducing cell and that proteins and DNA came along later. According to a model known as the RNA world, the chemistry of prebiotic Earth gave rise to self-replicating RNA molecules that functioned as both enzymes and substrates for their own replication. We represent the replication of RNA in the RNA world scenario as a circular arrow: RNA

(c) As the pre-cell system continues to evolve, its size and organization increase.

(d) The pre-cell system develops the ability to reproduce.

FIGURE 21-4 Origin of life: the metabolism first scenario

RNA often has catalytic properties; such enzymatic RNAs are called ribozymes. Before the evolution of true cells, ribozymes may have catalyzed their own replication in the clays, shallow rock pools, or hydrothermal vents where life originated. When RNA strands are added to a test tube containing RNA nucleotides but no enzymes, the nucleotides combine to form short RNA molecules. The occurrence of an RNA world early in the history of life can never be proven, but experiments with in vitro evolution, also called directed evolution, have shown it is feasible. These experiments address an important question about the RNA world, namely, could RNA molecules have catalyzed the many different chemical reactions needed for life? In directed evolution, a large pool of RNA molecules with different sequences is mixed, and molecules are selected for their ability to catalyze a single biologically important reaction (FIG. 21-5). Those molecules that have at least some catalytic ability are selected by researchers, who then amplify them into many copies. The researchers use certain chemicals to cause mutations, and then expose the molecules to another round of selection. After this cycle is repeated several times, the RNA molecules at the end of the selection process function efficiently as catalysts for the reaction. In vitro evolution studies have shown that RNA has a large functional repertoire—that is, RNA can catalyze a variety of biologically important reactions. Biologists hypothesize that in the RNA world, ribozymes initially catalyzed protein synthesis and other important biological reactions; only later did protein enzymes catalyze these reactions.

KEY EXPERIMENT QUESTION: Can in vitro natural selection result in chemical evolution of catalysts? HYPOTHESIS: Self-replicating RNA molecules can undergo changes that make them more efficient catalysts. EXPERIMENT: RNA molecules are selected from a large pool, based on their ability to catalyze a specific reaction, then amplified and mutated (see figure). This process is repeated 7 to 20 additional times.

Large pool of RNA molecules

Selection for ability to catalyze a chemical reaction

Molecules with some ability to catalyze the reaction

Amplification and mutation to create large pool of similar RNA molecules

Repeat the selection amplification mutation process

Molecules with best ability to catalyze the reaction

RESULTS AND CONCLUSION: The final group of RNA molecules is the most efficient at catalyzing the chemical reaction selected for. Scientists have developed more than two dozen synthetic RNA catalysts by in vitro evolution.

FIGURE 21-5 In vitro evolution of RNA molecules

RNA S protein

Interestingly, in test tube experiments, RNA molecules have evolved that direct protein synthesis by catalyzing peptide bond formation. Some single-stranded RNA molecules fold back on

themselves as a result of interactions among the nucleotides composing the RNA strand. Sometimes the conformation (shape) of the folded RNA molecule is such that it weakly binds to an amino acid (see discussion of peptidyl transferase in Chapter 13). Amino acids held close to one another by RNA molecules may bond, forming a polypeptide. We have considered how the evolution of informational molecules may have given rise to RNA and later to proteins. If a selfreplicating RNA capable of coding for proteins appeared prior to DNA, how did DNA, the universal molecule of heredity in cells, become involved? Perhaps RNA made double-stranded copies of itself that eventually evolved into DNA. DNA — RNA ¡ protein

The incorporation of DNA into the information transfer system was advantageous because the double helix of DNA is more stable (less reactive) than the single strand of RNA. Such stability in a molecule that stores genetic information would have provided a decided advantage in the prebiotic world (as it does today). Thus, in the DNA/RNA/protein world, DNA became the information storage molecule, RNA remained involved in protein synthesis, and protein enzymes catalyzed most cell reactions, including DNA replication and RNA synthesis. DNA S RNA S protein

RNA is still a necessary component of the information transfer system because DNA is not catalytic. Thus, natural selection at the molecular level favored the DNA ¡ RNA ¡ protein information sequence. Once DNA was incorporated into this sequence, RNA molecules assumed their present role as an intermediary in the transfer of genetic information. Several additional steps had to occur before a true living cell could evolve from macromolecular aggregations. For example, the self-replicating genetic code must have arisen extremely early in the prebiotic world because all organisms possess it—but how did it originate? Also, how did an encapsulating membrane of lipid and protein evolve? These kinds of questions remain for scientists to address.

Biological evolution began with the first cells No one knows exactly when or where the first cells appeared on Earth. Did they live at hydrothermal vents or at Earth’s surface? Were they bacteria? We cannot answer these questions, in part because no fossils exist that trace the transition from nonlife to life. Nonfossil evidence—isotopic “fingerprints” of organic carbon in ancient rocks in Greenland—suggest to some researchers that life existed as early as 3.8 billion years ago (bya). However, other scientists vigorously debate this conclusion. Microfossils, ancient remains of microscopic life, suggest that cells may have been thriving as long as 3.5 bya. Rich deposits of microfossils appear to exist in the Pilbara rocks of northwestern Australia and the Barberton rocks in South Africa; both of these deposits are about 3.3 billion to 3.5 billion years old. However, some scientists have challenged the interpretation of the oldest

Stanley M. Awramik/ Biological Photo Service

Fred Bavendam/ Peter Arnold, Inc.

(a) These living stromatolites at Hamlin Pool in Western Australia consist of mats of cyanobacteria (unicellular photosynthetic prokaryotes) and minerals such as calcium carbonate. They are several thousand years old.

(b) Cutaway view of a fossil stromatolite showing the layers of microorganisms and sediments that accumulated over time. This stromatolite, also from Western Australia, is about 3.5 billion years old.

FIGURE 21-6 Stromatolites

carbon-rich “squiggles” in ancient rocks as microfossils of prokaryotic organisms. They suggest that microfossils are not fossils at all but are formed by natural geologic processes. Evidence suggests that the earliest cells were prokaryotic. Stromatolites, another type of fossil evidence of the earliest cells, are rocklike columns composed of many minute layers of prokaryotic cells—that is, microbial biofilms (FIG. 21-6). Over time, sediment collects around the cells and mineralizes. Meanwhile, a new layer of living cells grows over the older, dead cells. Fossil stromatolite reefs are found in several places in the world, including Great Slave Lake in Canada and the Gunflint Iron Formations along Lake Superior in the United States. Some fossil stromatolites are extremely ancient. One group in Western Australia, for example, has been dated at more than 3 billion years. Living stromatolite reefs are rare but are found in certain hot springs and warm, shallow pools of fresh and salt water. Interestingly, in 2008 scientists reported that the microorganisms inhabiting living stromatolite communities are strikingly different from microorganisms in all other ecosystems studied to date; this discovery suggests that the microorganisms living in today’s stromatolites may have evolved from microorganisms living in the ancient stromatolite ecosystems.

The first cells were probably heterotrophic The earliest cells probably did not synthesize the organic molecules they needed but instead obtained them from the environment. These primitive heterotrophs may have consumed many types of organic molecules that had formed spontaneously—for example, sugars, nucleotides, and amino acids. By fermenting these organic compounds, they obtained the energy they needed to support life. Fermentation is, of course, an anaerobic process (performed in the absence of oxygen), and the first cells were almost certainly anaerobes.

When the supply of spontaneously generated organic molecules gradually declined, only certain organisms could survive. Mutations had probably already occurred that permitted some cells to obtain energy directly from sunlight, perhaps by using sunlight to make ATP. These cells, which did not require the energy-rich organic compounds that were now in short supply in the environment, had a distinct selective advantage. Photosynthesis requires both light energy and a source of electrons, which are used to reduce CO2 to form organic molecules such as glucose. (Recall the discussion of photosynthesis in Chapter 9.) Most likely, the first photosynthetic autotrophs—organisms that produce their own food from simple raw materials—used the energy of sunlight to split hydrogen-rich molecules such as H2S, releasing elemental sulfur (not oxygen) in the process. Splitting H2S is energetically much easier than splitting H2O because sulfur is not electronegative like oxygen (see Chapter 9). Indeed, the green sulfur bacteria and the purple sulfur bacteria still use H2S as a hydrogen (electron) source for photosynthesis. (Members of a third bacterial group, the purple nonsulfur bacteria, use other organic molecules or hydrogen gas as a hydrogen source.) The first photosynthetic autotrophs to obtain hydrogen electrons by splitting water were the cyanobacteria. Water was quite abundant on early Earth, as it is today, and the selective advantage of splitting water allowed cyanobacteria to thrive. The process of splitting water released oxygen as a gas (O2). Initially, the oxygen released during photosynthesis oxidized minerals in the ocean and in Earth’s crust, and oxygen did not begin to accumulate in the atmosphere for a long time. Eventually, however, oxygen levels increased in the ocean and the atmosphere. Scientists estimate the timing of the events just described on the basis of geologic and fossil evidence. Fossils from that period, which include rocks containing traces of chlorophyll, as well as the fossil stromatolites discussed earlier, indicate that the first photosynthetic organisms may have appeared as early as 3.5 bya.

This evidence suggests that heterotrophic forms may have existed even earlier.

KEY POINT

Ozone (O3) forms in the upper atmosphere when ultraviolet radiation from the sun breaks the double bonds of oxygen molecules.

Ultraviolet rays Upper atmosphere

3(O2)

2(O3) Lower atmosphere

Based on sulfur isotope data from ancient rocks in South Africa, it appears that cyanobacteria had produced enough oxygen to begin significantly changing the composition of the atmosphere by 2.4 bya. This date may be changed as more data accumulate: in 2009 geoscientists at Pennsylvania State University found strong isotopic evidence in Australia that oxygen was present in the oceans as early as 3.46 bya. Scientists are now looking for the same kind of evidence at other sedimentary layers known to be older than 2.4 bya. The increase in atmospheric oxygen affected life profoundly. The oxygen poisoned obligate anaerobes (organisms that cannot use oxygen for cellular respiration), and many species undoubtedly perished. Some anaerobes, however, survived in environments where oxygen did not penetrate; adaptations evolved in others that neutralized the oxygen so it could not harm them. In some organisms, called aerobes, a respiratory pathway evolved that used oxygen to extract more energy from food. Aerobic respiration was joined with the existing anaerobic process of glycolysis. The evolution of organisms that could use oxygen in their metabolism had several consequences. Organisms that respire aerobically gain much more energy from a single molecule of glucose than anaerobes gain by fermentation. (Recall the comparison of fermentation and aerobic respiration in Table 8.2.) As a result, the newly evolved aerobic organisms were more efficient and more competitive than anaerobes. Coupled with the poisonous nature of oxygen to many anaerobes, the efficiency of aerobes forced anaerobes into relatively minor roles. Today, the vast majority of organisms, including plants, animals, and most fungi, protists, archaea, and bacteria, use aerobic respiration; only a few archaea and bacteria and even fewer protists and fungi are anaerobic. The evolution of aerobic respiration stabilized both oxygen and carbon dioxide levels in the biosphere. Photosynthetic organisms used carbon dioxide as a source of carbon for synthesizing organic compounds. This raw material would have been depleted from the atmosphere in a relatively brief period without the advent of aerobic respiration, which released carbon dioxide as a waste product from the complete breakdown of organic molecules. Carbon thus started cycling in the biosphere, moving from the nonliving physical environment, to photosynthetic organisms, to heterotrophs that ate the photosynthetic organisms (see Chapter 55). Aerobic respiration released carbon back into the physical environment as carbon dioxide, and the carbon cycle continued. In a similar manner, molecular oxygen was produced by photosynthesis and used during aerobic respiration. Another significant consequence of photosynthesis occurred in the upper atmosphere, where molecular oxygen reacted to form ozone (O3) (FIG. 21-7). A layer of ozone eventually blanketed Earth, preventing much of the sun’s ultraviolet radiation from penetrating to the surface. With the ozone layer’s protection from the mutagenic effect of ultraviolet radiation, organisms could live closer to the surface in aquatic environments and eventually move

Sun

Aerobes appeared after oxygen increased in the atmosphere

FIGURE 21-7 The formation of ozone

onto land. Because the energy in ultraviolet radiation may have been necessary to form organic molecules, however, their abiotic synthesis decreased.

Eukaryotic cells descended from prokaryotic cells Eukaryotes may have appeared in the fossil record as early as 2.2 bya, and geochemical evidence suggests that eukaryotes were present much earlier. Steranes, molecules derived from steroids, have been discovered in Australian rocks dated at 2.7 billion years old. Because bacteria are not known to produce steroids, the steranes may be biomarkers for eukaryotes. These ancient rocks lack

fossil traces of ancient organisms because the rocks have since been exposed to heat and pressure that would have destroyed any ORIGINAL DNA fossilized cells. Steranes, however, are very PROKARYOTIC HOST CELL stable in the presence of heat and pressure. Eukaryotes arose from prokaryotes. Recall from Chapter 4 that archaeal and bacterial cells lack nuclear envelopes as well as Aerobic bacteria other membranous organelles such as mitoMultiple invaginations chondria and chloroplasts. How did these of the plasma membrane organelles arise? According to the hypothesis of serial endosymbiosis, organelles such as mitochondria and chloroplasts may have originated from mutually advantageous symbiotic relationships between two prokaryotic organisms (FIG. 21-8). ChloroEndoplasmic reticulum plasts apparently evolved from photosynAerobic bacteria and nuclear envelope become mitochondria thetic bacteria (cyanobacteria) that lived form from the plasma inside larger heterotrophic cells, whereas membrane invaginations (not part of serial mitochondria presumably evolved from aerendosymbiosis) obic bacteria (perhaps ancient purple bactePhotosynthetic ria) that lived inside larger anaerobic cells. bacteria become chloroplasts Thus, early eukaryotic cells were assemblages of formerly free-living prokaryotes. How did these bacteria come to be endosymbionts, which are organisms that live symbiotically inside a host cell? They may originally have been ingested, but not EUKARYOTIC EUKARYOTIC CELLS: CELLS: PLANTS, digested, by a host cell. Once incorporated, ANIMALS, FUNGI, SOME PROTISTS they could have survived and reproduced SOME PROTISTS along with the host cell so that future genFIGURE 21-8 Animated Serial endosymbiosis erations of the host also contained endosymbionts. A mutualistic relationship evolved between these two organisms in which each contributed mitochondria and chloroplasts. The outer membrane apparently something to the other. Eventually, the endosymbiont lost the developed from the invagination (infolding) of the host cell’s ability to exist outside its host, and the host cell lost the ability to plasma membrane, whereas the inner membrane is derived from survive without its endosymbionts. This hypothesis stipulates that the endosymbiont’s plasma membrane. (Serial endosymbiosis is each of these partners brought to the relationship something the discussed in greater detail in Chapter 26.) other lacked. For example, mitochondria provided the ability to Many endosymbiotic relationships exist today. Many corals have carry out the aerobic respiration lacking in the original anaerobic algae living as endosymbionts within their cells (see Fig. 54-12). host cell. Chloroplasts provided the ability to use a simple carbon Protozoa (Myxotricha paradoxa) live symbiotically in the gut of source (CO2) to produce needed organic molecules. The host cell termites; in turn, several different endosymbionts, including spiprovided endosymbionts with a safe environment and raw materirochete bacteria, attach to the protozoa and function as whiplike als or nutrients. flagella, allowing the protozoa to move. The principal evidence in favor of serial endosymbiosis is Although much evidence supports serial endosymbiosis, it that mitochondria and chloroplasts possess some (although not does not completely explain the evolution of eukaryotic cells from all) of their own genetic material and translational components. prokaryotes. For example, serial endosymbiosis does not currently They have their own DNA (as a circular molecule similar to that explain how a double membranous envelope came to surround the of archaea and bacteria; discussed in Chapter 25) and their own genetic material in the nucleus. ribosomes (which resemble bacterial rather than eukaryotic ribosomes). Mitochondria and chloroplasts also possess some of the Review machinery for protein synthesis, including tRNA molecules, and conduct protein synthesis on a limited scale independent of the ■ What major steps probably occurred in the origin of cells? nucleus. Furthermore, it is possible to poison mitochondria and ■ How did the presence of molecular oxygen in the atmosphere affect early life? chloroplasts with an antibiotic that affects bacteria but not eukary■ What kinds of evidence support serial endosymbiosis? otic cells. As discussed in Chapter 4, double membranes envelope

21.3 THE HISTORY OF LIFE ■ ■ LEARNING OBJECTIVE 6 Briefly describe the distinguishing organisms and major biological

The sequence of biological, climate, and geologic events that make up the history of life is recorded in rocks and fossils. The sediments of Earth’s crust consist of five major rock strata (layers), each subdivided into minor strata, lying one on top of the other. Very few places on Earth show all layers, but the strata present typically occur in the correct order, with younger rocks on top of older ones. These sheets of rock formed from the accumulation of mud and sand at the bottoms of the ocean, seas, and lakes. Each layer contains certain characteristic index fossils that identify deposits made at about the same time in different parts of the world. Geologists divide Earth’s history into units of time based on major geologic, climate, and biological events. Eons are the largest divisions of the geologic time scale. Eons are divided into eras; where fossil evidence exists, these divisions are based primarily on organisms that characterized each era. Eras are subdivided into periods, which in turn are composed of epochs. Relatively little is known about Earth from its beginnings approximately 4.6 bya up to 2.5 bya, a time known as the Archaean eon (TABLE 21-1). Life originated on Earth during the Archaean eon. Signs of life date as early as 3.5 bya. Not much physical evidence is available because the rocks of the Archaean eon, being extremely ancient, are deeply buried in most parts of the world. Ancient rocks are exposed in a few places, including the bottom of the Grand Canyon and along the shores of Lake Superior. Many Archaean rock formations have revealed what appear to be microfossils of cyanobacteria and other bacteria.

Rocks from the Ediacaran period contain fossils of cells and simple animals That part of time from 2500 mya to 542 mya is known as the Proterozoic eon. This enormous span of time is easier to study than the preceding Archaean eon because the rocks are less altered by heat and pressure. Life at the beginning of the Proterozoic eon consisted of prokaryotes such as cyanobacteria. Stromatolites were abundant. About 2.2 bya the first eukaryotic cells appeared. By the end of the Proterozoic eon, multicellularity was evident in the abundant fossils of small, soft-bodied, invertebrate animals. The Ediacaran period (pronounced “ee-dee-ack′-uh-run”), from 600 mya to 542 mya, is the last (most recent) period of the Proterozoic eon. It is named for the fossil deposits in the Ediacara Hills in South Australia, although Ediacaran fossils are also found in Newfoundland, the Mackenzie Mountains of northwestern Canada, the Doushantuo Formation of China, and other locations. Currently, more than 270 Ediacaran species have been identified and described. Ediacaran fossils are the oldest known fossils of multicellular animals (FIG. 21-9). Experts have not yet identified all the

Chase Studio/Photo Researchers, Inc.

events of the Ediacaran period and the Paleozoic, Mesozoic, and Cenozoic eras.

FIGURE 21-9 Reconstruction of life in an Ediacaran sea The organisms shown here are based on fossils from the Ediacara Hills of South Australia, although similar associations have been found in Ediacaran rocks from every continent except Antarctica. Shown are organisms that some scientists have interpreted as jellyfish, flatworms, algae, and paddleshaped organisms similar in appearance to soft corals.

simple, soft-bodied animals found in the Ediacara Hills and at other sites. Some paleontologists interpret many of these fossils as early sponges, jellyfish, corals, and comb jellies; however, other scientists think that the Ediacaran animals were not ancestral to modern-day species but instead became extinct at the end of the Ediacaran period.

A diversity of organisms evolved during the Paleozoic era The Paleozoic era began approximately 542 mya and lasted about 291 million years. It is divided into six periods: Cambrian, Ordovician, Silurian, Devonian, Carboniferous, and Permian. Rocks rich in fossils represent the oldest subdivision of the Paleozoic era, the Cambrian period. For about 40 million years, evolution was in such high gear, with the sudden appearance of many new animal body plans, that this period is called the Cambrian Radiation, or more informally, Cambrian explosion. Fossils of all contemporary animal phyla are present, along with many bizarre, extinct phyla, in marine sediments. The seafloor was covered with sponges, corals, sea lilies, sea stars, snails, clamlike bivalves, primitive squidlike cephalopods, lamp shells (brachiopods), and trilobites (see Fig. 31-26). In addition, small vertebrates—cartilaginous fishes—became established in the marine environment. Scientists have not determined the factors responsible for the Cambrian Radiation, a period unmatched in the evolutionary history of life. There is some evidence that oxygen concentrations, which had continued to gradually increase in the atmosphere, passed some critical threshold late in the Proterozoic eon. Scientists who advocate the oxygen enrichment hypothesis note that until late in the Proterozoic eon, Earth did not have enough oxygen to support larger animals. The most important fossil sites that document the Cambrian Radiation are the Chengjiang site in China (for Early Cambrian fossils) and the Burgess Shale in British Columbia (for Middle Cambrian fossils) (FIG. 21-10).

TABLE 21-1

Some Important Biological Events in Geologic Time

Eon

Period

Neogene period

Epoch Holocene epoch

Pleistocene epoch

Time* 0.01 (10,000 years ago)

2.6 mya

Miocene epoch

23 mya

Flowering plants continue to diversify; diversity of songbirds and grazing mammals

Oligocene epoch

34 mya

Spread of forests; apes appear; present mammalian families are represented

Eocene epoch

56 mya

Flowering plants dominant; modern mammalian orders appear and diversify; modern bird orders appear

Paleocene epoch

66 mya

Semitropical vegetation (flowering plants and conifers) widespread; primitive mammals diversify

146 mya

Rise of flowering plants; dinosaurs reach peak, then become extinct at end; toothed birds become extinct

200 mya

Gymnosperms common; large dinosaurs; first toothed birds

251 mya

Gymnosperms dominant; ferns common; first dinosaurs; first mammals

299 mya

Conifers diversify; cycads appear; modern insects appear; mammal-like reptiles; extinction of many invertebrates and vertebrates at end of Permian

359 mya

Forests of ferns, club mosses, horsetails, and gymnosperms; many insect forms; spread of ancient amphibians; first reptiles

416 mya

First forests; gymnosperms appear; many trilobites; wingless insects appear; fishes with jaws appear and diversify; amphibians appear

444 mya

Vascular plants appear; coral reefs common; jawless fishes diversify; terrestrial arthropods

488 mya

Fossil spores of terrestrial plants (bryophytes?); invertebrates dominant; coral reefs appear; first fishes appear

542 mya

Bacteria and cyanobacteria; algae; fungi; age of marine invertebrates; first chordates

600 mya

Algae and soft-bodied invertebrates diversify

Jurassic period

Carboniferous period

Paleozoic era

Extinction of some plant species; extinction of many large mammals at end Expansion of grasslands and deserts; many grazing animals

Permian period

Devonian period

Silurian period

Ordovician period

Cambrian period

Proterozoic eon

Decline of some woody plants; rise of herbaceous plants; age of Homo sapiens

5 mya

Triassic period

Archaean eon

Some Important Biological Events

Pliocene epoch

Cretaceous period Mesozoic era

Phanerozoic eon

Paleogene period

Cenozoic era

Quaternary period

Era

Ediacaran period

Early Proterozoic period

Eukaryotes evolve 2500 mya Oldest known rocks; prokaryotes evolve; atmospheric oxygen begins to increase 4600 mya

*Time from beginning of eon, period, or epoch to present (millions of years).

Both: National Museum of Natural History, Courtesy of Chip Clark © 2006 Smithsonian Institute

bony fishes: lobe-finned fishes and rayfinned fishes, which gave rise to the major orders of modern fishes (see Chapter 31). Tiktaalik was a transitional form between fishes and tetrapods, which are vertebrates with four limbs (see Fig. 32-17). Tiktaalik is considered a fish because it had scales and fins. Like tetrapods, however, Tiktaalik had a movable neck and ribs that enclosed lungs. Upper (more recent) Devonian sediments contain fossil remains of salamander-like amphibians (labyrin(a) Marrella splendens was a small (b) Wiwaxia was a bristle-covered marine arthropod with 26 body segments. It is the worm that was distantly related to thodonts) that were often quite large, with most abundant fossil arthropod found in the earthworms. It had scaly armor and short necks and heavy, muscular tails. Burgess Shale. needlelike spines for protection. Wingless insects also originated in the Late Devonian period. FIGURE 21-10 Fossils from the Cambrian Radiation The early vascular plants (plants with These fossils were discovered in the Burgess Shale in the Canadian Rockies of British Columbia. specialized tissue to conduct water and nutrients) diversified during the Devonian According to geologists, seas gradually flooded the continents period in a burst of evolution that rivaled that of animals during during the Cambrian period. In the Ordovician period, much the Cambrian Radiation. With the exception of flowering plants, land was covered by shallow seas, in which there was another burst all major plant groups appeared during the Devonian period. Forof evolutionary diversification, although not as dramatic as the ests of ferns, club mosses, horsetails, and seed ferns (an extinct Cambrian Radiation. The Ordovician seas were inhabited by giant group of ancient plants that had fernlike foliage but reproduced by cephalopods, squidlike animals with straight shells 5 to 7 m (16 to forming seeds) flourished. 23 ft) long and 30 cm (12 in) in diameter. Coral reefs first appeared The Carboniferous period is named for the great swamp forduring this period, as did small, jawless, bony-armored fishes called ests whose remains persist today as major coal deposits. Much of ostracoderms (FIG. 21-11). Lacking jaws, these fishes typically had the land during this time was covered with low swamps filled with round or slitlike mouth openings that may have sucked in small horsetails, club mosses, ferns, seed ferns, and gymnosperms, which food particles from the water or scooped up organic debris from are seed-bearing plants such as conifers (FIG. 21-12). Amphibians, which underwent an adaptive radiation and exploited both aquatic the bottom. Ordovician deposits also contain fossil spores of terand terrestrial ecosystems, were the dominant terrestrial carnivores restrial (land-dwelling) plants, suggesting the colonization of land of the Carboniferous period. Reptiles first appeared and diverged to had begun. form two major lines at this time. One line consisted of mostly small During the Silurian period, jawless fishes diversified considand mid-sized insectivorous (insect-eating) lizards; this line later erably, and jawed fishes first appeared. Definitive evidence of two led to lizards, snakes, crocodiles, dinosaurs, and birds. The other life-forms of great biological significance appeared in the Silurian reptilian line led to a diverse group of Permian and Early Mesozoic period: terrestrial plants and air-breathing animals. The evolumammal-like reptiles. Two groups of winged insects, cockroaches tion of plants allowed animals to colonize the land because plants and dragonflies, appeared in the Carboniferous period. provided the first terrestrial animals with food and shelter. All airbreathing land animals discovered in Silurian rocks were arthropods—millipedes, spiderlike (b) Pterapsis arthropods, and possibly centipedes. From an ecological perspective, the energy flow from plants to animals probably occurred via detritus, which is organic debris from decomposing organisms, rather than directly from living plant material. Millipedes eat plant detritus today, and spiders and centipedes prey on millipedes and other animals. The Devonian period is frequently called (c) Jamoytius the Age of Fishes. This period witnessed the explosive radiation of fishes with jaws, an adaptation that lets a vertebrate chew and bite. (a) Thelodus Armored placoderms, an extinct group of jawed fishes, diversified to exploit varied lifestyles (see FIGURE 21-11 Ostracoderms Fig. 32-10). Appearing in Devonian deposits Ostracoderms, primitive jawless fishes that lived in the Ordovician, Silurian, and Devonian periods, ranged from 10 to 50 cm (4 to 20 in) in length. are sharks and the two predominant types of

Field Museum of Natural History, Chicago, No. GEO85638c

FIGURE 21-12 Reconstruction of a Carboniferous forest Plants of this period included giant ferns, horsetails, club mosses, seed ferns, and early gymnosperms.

Amphibians continued in importance during the Permian period, but they were no longer the dominant carnivores in terrestrial ecosystems. During the Permian period, reptiles diversified and dominated both carnivorous and herbivorous terrestrial lifestyles. One important group of mammal-like reptiles, originating in the Permian and extending into the Mesozoic era, were the therapsids, a group that included the ancestor of mammals (see Fig. 32-24). During the Permian period, seed plants diversified and dominated most plant communities. Cone-bearing conifers were widespread, and cycads (plants with crowns of fernlike leaves and large, seedcontaining cones) and ginkgoes (trees with fan-shaped leaves and exposed, fleshy seeds) appeared. The greatest mass extinction of all time occurred at the end of the Paleozoic era, between the Permian and Triassic periods, 251 mya. More than 90% of all existing marine species became extinct at this time, as did more than 70% of the vertebrate genera living on land. There is also evidence of a major extinction of plants. Many causes for the Late Permian mass extinction have been suggested, from meteor impacts to global warming to changes in ocean chemistry. Regardless of cause, evidence suggests that the extinction occurred globally in a very compressed period, within a few hundred thousand years. This is an extremely short period in the geologic time scale and suggests that some sort of catastrophic event caused the mass extinction.

Dinosaurs and other reptiles dominated the Mesozoic era The Mesozoic era began about 251 mya and lasted some 185 million years. It is divided into the Triassic, Jurassic, and Cretaceous periods. Fossil deposits from the Mesozoic era occur worldwide.

Notable sites include the Yixian Formation in northeastern China, the Solnhofen Limestone in Germany, northwestern Patagonia in Argentina, the Sahara in central Niger, the Badlands in South Dakota, and other sites in western North America. The outstanding feature of the Mesozoic era was the origin, differentiation, and ultimately the extinction of a large variety of reptiles. For this reason, the Mesozoic era is commonly called the Age of Reptiles. Most of the modern orders of insects appeared during the Mesozoic era. Snails and bivalves (clams and their relatives) increased in number and diversity, and sea urchins reached their peak diversity. From a botanical viewpoint, the Mesozoic era was dominated by gymnosperms until the Mid-Cretaceous period, when the flowering plants first diversified. During the Triassic period, reptiles underwent an adaptive radiation leading to the formation of many groups. On land, the dominant Triassic groups were the mammal-like therapsids, which ranged from small-sized insectivores (insect-eating reptiles) to moderately large herbivores (plant-eating reptiles), and a diverse group of thecodonts, early “ruling reptiles,” that were primarily carnivores. Thecodonts are the ancestral reptiles that gave rise to crocodilians, flying reptiles, dinosaurs, and birds. In the ocean, several important marine reptile groups, the plesiosaurs and ichthyosaurs, appeared in the Triassic period. Plesiosaurs had bodies up to 15 m (about 49 ft) long and paddlelike fins. Ichthyosaurs had body forms superficially resembling those of sharks or porpoises, with short necks, large dorsal fins, and sharktype tails (FIG. 21-13). Ichthyosaurs had very large eyes, which may have helped them see at diving depths of 500 m (1650 ft) or more. Pterosaurs, the first flying reptiles, appeared and underwent considerable diversification during the Mesozoic era (FIG. 21-14). This group produced some quite spectacular forms, most notably

© John Sibbick

© John Sibbick

FIGURE 21-14 Pterosaurs

Brachypterygius (also known by the genus Grendelius) was an ichthyosaur that superficially resembled a shark or porpoise. It had large teeth and strong jaws and was about 4 m (13 ft) long.

Shown are Peteinosaurus (left), with a wingspan of 60 cm (2 ft), and Eudimorphodon (right foreground), with a wingspan of 75 cm (2.5 ft). Both species had long, sharp teeth for catching insects or fishes while flying.

the giant Quetzalcoatlus, known from fragmentary Cretaceous fossils in Texas to have had a wingspan of 11 to 15 m (36 to 49 ft). Pterosaur wings were leathery membranes of skin that were supported by an elongated fourth finger bone. Claws extended from the other finger bones. The first mammals to appear in the Triassic period were small insectivores that evolved from the mammal-like therapsids. Mam-

mals diversified into a variety of mostly small, nocturnal insectivores during the remainder of the Mesozoic era, with marsupial and placental mammals appearing later in the Mesozoic. During the Jurassic and Cretaceous periods, crocodiles, lizards, snakes, and birds appeared, and the dinosaurs diversified dramatically (FIG. 21-15). The mosasaurs, one group of lizards, were large, voracious marine predators during the Late Cretaceous

© John Sibbick

FIGURE 21-13 Ichthyosaurs

FIGURE 21-15 Dinosaurs Three Deinonychus dinosaurs attack a larger Tenontosaurus. The name Deinonychus means “terrible claw” and refers to the enlarged, sharp claw on the second digit of its hind feet. Deinonychus dinosaurs were small (3 m, or 10 ft in length) but fearsome predators that hunted in packs. Tenontosaurus adults were as long as 7.5 m (24 ft).

ducklike, hence the common name duck-billed dinosaurs. Other ornithischians had great armor plates, possibly as protection against carnivorous saurischians. Ankylosaurus, for example, had a broad, flat body covered with armor plates (actually bony scales embedded in the skin) and large, laterally projecting spikes. Over the past few decades, scientists have reconsidered many traditional ideas about dinosaurs and no longer think they were all cold-blooded, slow-moving monsters living in swamps. Recent evidence suggests that at least some dinosaurs were warm-blooded, agile, and able to move extremely fast. Many dinosaurs appear to have had complex social behaviors, including courtship rituals and parental nurturing of their young. Some species lived in social groups and hunted in packs. Birds appeared by the Late Jurassic period, and fossil evidence indicates they evolved directly from saurischian dinosaurs (see Inquiring About: The Origin of Flight in Birds). Archaeopteryx, the oldest known bird in the fossil record, lived about 150 mya (see Fig. 32-22b). It was about the size of a pigeon and had rather feeble wings that it used to glide rather than actively fly. Although Archaeopteryx is considered a bird, it had many reptilian features, including a mouthful of teeth and a long, bony tail. Thousands of well-preserved Saurischians bird fossils have been found in Ilium Early Cretaceous deposits in Hip socket China. These include Sinornis, a Pubis 135-million-year-old sparrow-sized bird capable of perching, and the magpie-sized Confuciusornis, the Ischium earliest known bird with a toothCoelophysis less beak. Confuciusornis may date back as far as 142 mya. At the end of the Cretaceous period, 66 mya, dinosaurs, pterosaurs, and many other animals (a) The saurischian pelvis. Note the opening (hip socket), a trait possessed by no abruptly became extinct. Many quadrupedal vertebrates other than dinosaurs. gymnosperms, with the exception of conifers, also perished. Evidence suggests that a catastrophic collision of a large extraterrestrial Ornithischians body with Earth dramatically Ilium changed the climate at the end of the Cretaceous period. Part of Hip socket the evidence is a thin band of dark clay, with a high concentration of iridium, located between MesoPubis zoic and Cenozoic sediments at Ischium more than 200 sites around the world. Iridium is rare on Earth but abundant in meteorites. The force Stegosaurus of the impact would have driven the iridium into the atmosphere, (b) The ornithischian pelvis. Note that it has the hole in the hip socket but differs from the saurischian pelvis in that it has a backward-directed extension of the pubis. to be deposited later on the land by precipitation. FIGURE 21-16 Saurischian and ornithischian dinosaurs The Chicxulub crater, buried In each dinosaur figure, the pale yellow femur is shown relative to the pelvic bone. under the Yucatán Peninsula in period. The mosasaurs, which are now extinct, attained lengths of 10 m (33 ft) or more. The evolutionary radiation of the dinosaurs expanded from one lineage to several dozen that ecologically filled a variety of adaptive zones. Dinosaurs are placed in two main groups based on their pelvic bone structure: the saurischians and the ornithischians (FIG. 21-16). Some saurischians were fast, bipedal (walking on two feet) forms ranging from those the size of a dog to the ultimate representatives of this group, the gigantic carnivores of the Cretaceous period—Tyrannosaurus, Giganotosaurus, and Carcharodontosaurus. Other saurischians were huge, quadrupedal (walking on four feet) dinosaurs that ate plants. Some of these were the largest terrestrial animals that have ever lived, including Argentinosaurus, with an estimated length of 30 m (98 ft) and an estimated weight of 72 to 90 metric tons (80 to 100 tons). The other group of dinosaurs, the ornithischians, was entirely herbivorous. Although some ornithischians were bipedal, most were quadrupedal. Some had no front teeth and possessed stout, horny, birdlike beaks. In some species these beaks were broad and

In q u ir in g A b o u t THE ORIGIN OF FLIGHT IN BIRDS

1

Beebe, W. H. “A Tetrapteryx Stage in the Ancestry of Birds.” Zoologica, Vol. 2, 1915.

2 Xu, X., Z. Zhou, X. Wang, X. Kuang, F. Zhang, and X. Du. “Four-Winged Dinosaurs from China.” Nature, Vol. 421, Jan. 23, 2003.

maeosaur dinosaur were found in Liaoning Province in northeastern China. The dinosaur, Microraptor gui, had feathers on both forelimbs and hind limbs as well as on its long tail (see figure). It was small—77 cm (about 30 inches) in length, including the tail—and appeared to be adapted to life in the trees. The feathers on M. gui were similar to those of modern-day birds. Downy feathers covered the body, and each limb had about 12 “primary” flight feathers and about 18 shorter, “secondary” feathers. Microraptor gui’s flight feathers were asymmetrical, a characteristic associated with flight or gliding in modern birds. The primary and secondary feathers followed a similar pattern on both the forelimbs and hind limbs, and this pattern resembles that on modern birds. In part because M. gui’s breastbone was not structured to attach large flight muscles, the dinosaur probably glided rather than flapped its wings. Microraptor gui, which is about 126 million years old, is not a direct ancestor of birds. Birds had already evolved when M. gui existed. The earliest known bird, Archaeopteryx, lived about 150 mya and therefore predates M. gui by about 25 million years. Microraptor gui is considered to have evolved from a basal member—that is, an earlier evolutionary branch—of the most recent ancestor of Archaeopteryx and other birds. Like M. gui, earlier feathered dino-

Mexico, is the apparent site of this collision. The impact produced giant tsunamis (tidal waves) that deposited materials from the extraterrestrial body around the perimeter of the Gulf of Mexico, from Alabama to Guatemala. It may have caused global forest fires and giant smoke and dust clouds that lowered global temperatures for many years. Although scientists widely accept that a collision with an extraterrestrial body occurred 66 mya, they have reached no consensus about the effects of such an impact on organisms. The extinction of many marine organisms at or immediately after the time of the impact was probably the result of the environmental upheaval that the collision produced. However, many clam species associated with the mass extinction at the end of the Cretaceous period seem to have gone extinct before the impact, suggesting that other factors caused some of the massive extinctions occurring then.

Portia Sloan

Do we know how birds evolved? The evolution of birds is arguably one of the most interesting chapters in Earth’s history of life. Given the substantial fossil evidence, most paleontologists have concluded that the ancestors of birds were dinosaurs, specifically the dromaeosaurs, a group of grounddwelling, bipedal theropods (carnivorous saurischians). Beginning in 1997, paleontologists made several discoveries of fossil dinosaurs with feathers, indicating that feathers appeared before birds. Scientists think that feathers evolved as one or a series of evolutionary novelties, or preadaptations (see Chapter 20). The first feathers may have provided thermal insulation but were subsequently modified for flight. Once dinosaurs and early birds had feathers, how did they fly? Did tree-dwelling animals glide as an intermediate step in bird flight, or did ground-dwelling animals flap their wings and run to provide thrust and lift for a takeoff? The question about how flight originated in birds has intrigued biologists for more than a century. In 1915, U.S. zoologist William Beebe hypothesized that the ancestors of birds were probably tree-dwelling gliders that had feathers on all four limbs.1 Because no fossil evidence supported his suggestion, scientists did not consider it seriously at the time. Almost a century later, in 2003, a group of Chinese paleontologists announced the discovery of two nearly complete fossils of the organisms that Beebe had hypothesized.2 The fossils of a small, feathered dro-

saurs may have been four-winged organisms that lived and glided in the trees. During the course of bird evolution, the feathered hind limbs may have become reduced and eventually lost. However, an alternative hypothesis is that feathered hind limbs may have been a failed evolutionary experiment restricted to dromaeosaurs and not important as an intermediate step in bird evolution. Further analysis of M. gui and future discoveries of earlier dromaeosaur fossils may shed some light on the importance of feathered hind limbs. Microraptor gui has given paleontologists and biologists much to consider in the evolutionary transition from dinosaurs to birds. Scientists will continue to study and debate the evolution of flight in birds for many years.

The Cenozoic era is the age of mammals With equal justice the Cenozoic era could be called the Age of Mammals, the Age of Birds, the Age of Insects, or the Age of Flowering Plants. This era is marked by the appearance of all these forms in great variety and numbers of species. The Cenozoic era extends from 66 mya to the present. It is subdivided into three periods: the Paleogene period, encompassing some 43 million years, the Neogene period, which was 20.4 million years in length, and the Quaternary period, which covers the last 2.6 million years. The Paleogene period is subdivided into three epochs, named from earliest to latest: Paleocene, Eocene, and Oligocene. The Neogene period is subdivided into two epochs: the Miocene and Pliocene. The Quaternary period is subdivided into the Pleistocene and Holocene epochs.

© John Sibbick

© John Sibbick

Flowering plants, which arose during the Cretaceous period, continued to diversify during the Cenozoic era. During the Paleocene and Eocene epochs, fossils indicate that tropical and semitropical plant communities extended to relatively high latitudes. Palms, for example, are found in Eocene deposits in Wyoming. Later in the Cenozoic era, there is evidence of more open habitats. Grasslands and savannas spread throughout much of North America during the Miocene epoch, with deserts developing later in the Pliocene and Pleistocene epochs. During the Pleistocene epoch, plant communities changed dynamically in response to the fluctuating climates associated with the multiple advances and retreats of continental glaciers. During the Eocene epoch, there was an explosive radiation of birds, which acquired adaptations for different habitats. Paleontologists hypothesize that the jaws and beak of the flightless giant bird Diatryma, for example, may have been adapted primarily for crushing and slicing vegetation in Eocene forests, marshes, and grasslands. Other paleontologists hypothesize that these giant birds were carnivores that killed or scavenged mammals and other vertebrates (FIG. 21-17). During the Paleocene epoch, an explosive radiation of primitive mammals occurred. Most of these were small forest dwellers that are not closely related to modern mammals. During the Eocene epoch, mammals continued to diverge, and all the modern orders first appeared. Again, many of the mammals were small, but there were also some larger herbivores. During the Oligocene epoch, many modern families of mammals evolved, including the first apes in Africa. Many lineages showed adaptations that suggest a more open type of habitat, such as grassland or savanna. Many mammals were larger and had longer legs for running, specialized teeth for chewing coarse vegetation or for preying on animals, and increases in their relative brain

FIGURE 21-18 A mammal from the Oligocene epoch Paraceratherium was an indricothere, a hornless relative of the rhinoceros. This huge land mammal was about 8 m (26 ft) long and weighed about 15 to 20 tons. It probably ate leaves and branches of deciduous trees, much as a modern-day giraffe does.

sizes. The indricotheres, for example, are extinct relatives of the rhinoceros. These mammals, which lived on the grassless plains of Eurasia, became progressively larger during the Oligocene epoch (FIG. 21-18). Human ancestors appeared in Africa during the Late Miocene and Early Pliocene epochs. Homo, the genus to which humans belong, appeared approximately 2.5 mya. (Primate evolution, including human evolution, is discussed in Chapter 22.) The Pliocene and Pleistocene epochs witnessed the introduction of spectacular North and South American large-mammal fauna, including mastodons, saber-toothed cats, camels, giant ground sloths, and giant armadillos. However, many of the large mammals became extinct at the end of the Pleistocene epoch. This extinction was possibly due to climate change—the Pleistocene epoch was marked by several ice ages—or to the influence of humans, which had spread from Africa to Europe and Asia, and later to North and South America by crossing a land bridge between Siberia and Alaska. Archaeological evidence indicates that this mass extinction event was concurrent with the appearance of human hunters.

Review ■

FIGURE 21-17 A bird from the Eocene epoch

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The flightless bird Diatryma, which stood 2.1 m (7 ft) tall and weighed about 175 kg (385 lb), may have been an herbivore or a formidable predator. In this picture, Diatryma has captured a small, horselike perissodactyl.

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What is the correct order of appearance in the fossil record, starting with the earliest: eukaryotic cells, multicellular organisms, prokaryotic cells? What is the correct order of appearance in the fossil record, starting with the earliest: reptiles, mammals, amphibians, fishes? What is the correct order of appearance in the fossil record, starting with the earliest: flowering plants, ferns, gymnosperms?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

21.1 (page 447) 1 Describe the conditions that scientists think existed on early Earth. ■

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Biologists generally agree that life originated from nonliving matter by chemical evolution. Hypotheses about chemical evolution are testable. Four requirements for chemical evolution are (1) the absence of oxygen, which would have reacted with and oxidized abiotically produced organic molecules; (2) energy to form organic molecules; (3) chemical building blocks, including water, minerals, and gases present in the atmosphere; and (4) sufficient time for molecules to accumulate and react. Learn more about conditions on early Earth by clicking on the figure in CengageNOW.

2 Contrast the prebiotic soup hypothesis and the iron–sulfur world hypothesis. During chemical evolution, small organic molecules formed spontaneously and accumulated. The prebiotic soup hypothesis proposes that organic molecules formed near Earth’s surface in a “sea of organic soup” or on rock or clay surfaces. The iron–sulfur world hypothesis suggests that organic molecules were produced at hydrothermal vents, cracks in the deep-ocean floor.

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See the Miller–Urey experiment unfold by clicking on the figure in CengageNOW.

21.2 (page 449) 3 Outline the major steps hypothesized to have occurred in the origin of cells. After small organic molecules formed and accumulated, macromolecules assembled from the small organic molecules. Macromolecular assemblages called protobionts formed from macromolecules. Cells arose from the protobionts. ■ According to a model known as the RNA world, RNA was the first informational molecule to evolve in the progression toward a selfsustaining, self-reproducing cell. Natural selection at the molecular level eventually resulted in the information sequence DNA ¡ RNA ¡ protein. 4 Explain how the evolution of photosynthetic autotrophs affected both the atmosphere and other organisms. ■ The first cells were prokaryotic heterotrophs that obtained organic molecules from the environment. They were almost certainly anaerobes. Later, autotrophs—organisms that produce their own organic molecules by photosynthesis—evolved. ■ The evolution of oxygen-generating photosynthesis ultimately changed early life. The accumulation of molecular oxygen in the atmosphere permitted the evolution of aerobes, organisms that could use oxygen for a more efficient type of cellular respiration. ■

5 Describe the hypothesis of serial endosymbiosis. ■

Eukaryotic cells arose from prokaryotic cells. According to the hypothesis of serial endosymbiosis, certain eukaryotic organelles (mitochondria and chloroplasts) evolved from prokaryotic endosymbionts incorporated within larger prokaryotic hosts. Learn more about serial endosymbiosis by clicking on the figure in CengageNOW.

21.3 (page 455) 6 Briefly describe the distinguishing organisms and major biological events of the Ediacaran period and the Paleozoic, Mesozoic, and Cenozoic eras. ■ Life at the beginning of the Proterozoic eon (2500 mya to 542 mya) consisted of prokaryotes. About 2.2 bya the first eukaryotic cells appeared. The Ediacaran period, from 600 mya to 542 mya, is the last period of the Proterozoic eon. Ediacaran fossils are the oldest known fossils of multicellular animals. Ediacaran fauna were small, soft-bodied invertebrates. During the Paleozoic era, which began about 542 mya and lasted ■ approximately 291 million years, all major groups of plants, except flowering plants, and all animal phyla appeared. Fishes and amphibians flourished, and reptiles appeared. The greatest mass extinction of all time occurred at the end of the Paleozoic era, 251 mya. More than 90% of marine species and 70% of land-dwelling vertebrate genera became extinct, as well as many plant species. ■ The Mesozoic era began about 251 mya and lasted some 185 million years. Flowering plants appeared, and reptiles diversified. Dinosaurs, which descended from early reptiles, dominated. Insects flourished, and birds and early mammals appeared. At the end of the Cretaceous period, 66 mya, many species abruptly became extinct. A collision of a large extraterrestrial body with Earth may have resulted in dramatic climate changes that played a role in this mass extinction.

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In the Cenozoic era, which extends from 66 mya to the present, flowering plants, birds, insects, and mammals diversified greatly. Human ancestors appeared in Africa during the Late Miocene and Early Pliocene epochs.

T E S T YO U R U N D E R S TA N D I N G 1. Energy, the absence of oxygen, chemical building blocks, and time were the requirements for (a) chemical evolution (b) biological evolution (c) the Cambrian Radiation (d) the mass extinction episode at the end of the Cretaceous period (e) directed evolution

2. Many scientists think that _____ was the first information molecule to evolve. (a) DNA (b) RNA (c) a protein (d) an amino acid (e) a lipid 3. The first cells were probably (a) heterotrophs (b) autotrophs (c) anaerobes (d) a and c (e) b and c

4. According to the hypothesis of serial endosymbiosis, (a) life originated from nonliving matter (b) the pace of evolution quickened at the start of the Cambrian period (c) chloroplasts, mitochondria, and possibly other organelles originated from intimate relationships among prokaryotic cells (d) banded iron formations reflect the buildup of sufficient oxygen in the atmosphere to oxidize iron at Earth’s surface (e) the first photosynthetic organisms appeared 3.1 bya to 3.5 bya 5. During the Early _____, life consisted of prokaryotic cells, but by the end of this geologic time span, multicellular eukaryotic organisms had evolved. (a) Cenozoic era (b) Paleozoic era (c) Mesozoic era (d) Archaean eon (e) Proterozoic eon 6. Geologists divide the eons into (a) periods (b) epochs (c) eras (d) millennia (e) none of the preceding 7. Ediacaran fossils (a) are the oldest known fossils of multicellular animals (b) come from the Burgess Shale in British Columbia (c) contain remains of large salamander-like organisms (d) are the oldest fossils of early vascular plants (e) contain a high concentration of iridium

period (c) Silurian period (d) Carboniferous period (e) Permian period 9. The greatest mass extinction episode in the history of life occurred at what boundary? (a) Pliocene–Pleistocene (b) Permian– Triassic (c) Mesozoic–Cenozoic (d) Cambrian–Ordovician (e) Triassic–Jurassic 10. The Age of Reptiles corresponds to the (a) Paleozoic era (b) Mesozoic era (c) Cenozoic era (d) Pleistocene epoch (e) Permian period 11. Evidence exists that a catastrophic collision between Earth and a large extraterrestrial body occurred 66 mya, resulting in the extinction of (a) worms, mollusks, and soft-bodied arthropods (b) jawless ostracoderms and jawed placoderms (c) dinosaurs, pterosaurs, and many gymnosperm species (d) mastodons, sabertoothed cats, and giant ground sloths (e) ferns, horsetails, and club mosses 12. Flowering plants and mammals diversified and became dominant during the (a) Paleozoic era (b) Mesozoic era (c) Cenozoic era (d) Devonian period (e) Cambrian period

8. The time of greatest evolutionary diversification in the history of life occurred during the (a) Cambrian period (b) Ordovician

CRITICAL THINKING

4. ANALYZING DATA. Evidence for the oldest cyanobacterial fossils (stromatolites) dates to about 2 billion years, but molecular evidence inferred from lipids in ancient rocks puts the date for living cyanobacteria and eukaryotes at 2.7 billion years. Develop two opposing hypotheses to explain the discrepancy between the dates.

2.0

Oldest structural fossils of cyanobacteria

2.4 2.5

Rise of atmospheric oxygen

2.7

Molecular fossils of cyanobacteria and eukaryotes

4.6

Archaean

3. EVOLUTION LINK. Why did the evolution of multicellular organisms such as plants and animals have to be preceded by the evolution of oxygen-producing photosynthesis?

Time (billions of years ago)

2. EVOLUTION LINK. If living cells were produced in a test tube from nonbiological components by chemical processes, would this accomplishment prove that life evolved in a similar manner billions of years ago? Why or why not?

Proterozoic

0.5

1. EVOLUTION LINK. If you were studying how protobionts evolved into cells and you developed a protobiont that was capable of self-replication, would you consider it a living cell? Why or why not?

Origin of Earth

Additional questions are available in CengageNOW at www.cengage.com/ login.

22

The Evolution of Primates

Des Bartlett/Photo Researchers, Inc.

Studying human evolution. The late Drs. Mary and Louis Leakey studied fossil teeth from Australopithecus boisei, an early hominin (human) that lived in Africa. This photo was taken in Olduvai Gorge in Tanzania. The Leakey family has contributed much of what we know about human evolution. Son Richard, daughter-in-law Meave, and granddaughter Louise have followed in the footsteps of Mary and Louis.

KEY CONCEPTS

T

welve years after Darwin wrote On the Origin of Species by Natural Selection, he published another controversial book, The Descent of

22.1 Humans are classified in the order Primates, along with lemurs, tarsiers, monkeys, and apes. This classification is based on close evolutionary ties.

Man, which addressed human evolution. In this book, Darwin hypothe-

22.2 The study of living primates provides clues to help

fairly incomplete. However, research during the last several decades,

scientists reconstruct the adaptations and lifestyles of early primates, some of which were ancestors of humans.

22.3 Fossil evidence indicates that the earliest human ancestors arose in Africa and shared many features with their apelike ancestors. The human brain did not begin to enlarge to its present size and complexity until long after human ancestors had evolved bipedal locomotion.

22.4 Human culture began when human ancestors started making stone tools.

sized that humans and apes share a common ancestry. For nearly a century after Darwin’s studies, fossil evidence of human ancestry remained especially in Africa, has yielded a rapidly accumulating set of fossils that gives an increasingly clear answer to the question, “Where did we come from?” (see photograph). Fossil evidence from paleoanthropology, the study of human evolution, has allowed scientists to infer not only the structure but also the habits of early humans and other primates. Teeth and bones are the main fossil evidence that paleoanthropologists study. Much information can be obtained by studying teeth, which have changed dramatically during

the course of primate and human evolution. Because tooth enamel is more mineralized (harder) than bone, teeth are more likely to fossilize. The teeth of each primate species, living or extinct, are distinctive enough to identify the species, approximate age, diet, and even sex of the individual. Based on fossil evidence, paleontologists hypothesize that the first primates descended from small, shrewlike placental mammals that lived in trees and ate insects. (Placental mammals are the largest and most successful group of mammals. They have a placenta, an organ that exchanges materials between the mother and the embryo/fetus developing in the uterus.) Many traits of the 233 living primate species are related to their arboreal (tree-dwelling)

they have stereoscopic (three-dimensional) vision, which is important in judging distance and in depth perception. Stereoscopic vision is vital in an arboreal environment, especially for species that leap from branch to branch, because an error in depth perception may cause a fatal fall. In addition to having sharp sight, primates hear acutely. Primates share several other characteristics, including a relatively large brain. Biologists have suggested that the increased sensory input associated with primates’ sharp vision and greater agility favored the evolution of larger brains. Primates are generally very social and intelligent animals that reach sexual maturity relatively late in life. They typically have long lifespans. Females usually bear one offspring at a time; the baby is helpless and requires a long period of nurturing and protection.

past. This chapter focuses on humans and their ancestors, who differ from most other primates because they did not remain in the

Review

trees but instead adapted to a terrestrial way of life. ■ ■

How are primate hands and feet adapted to an arboreal existence? Why is the location of primate eyes in the front of the head an important adaptation for an arboreal life?

22.1 PRIMATE ADAPTATIONS ■ ■ LEARNING OBJECTIVE 1

Describe the structural adaptations that primates have for life in treetops.

Humans and other primates—including lemurs, tarsiers, monkeys, and apes—are mammals that share such traits as flexible hands and feet with five digits, a strong social organization, and front-facing eyes, which permit depth perception. Mammals (class Mammalia) evolved from mammal-like reptiles known as therapsids more than 200 million years ago (mya), during the Mesozoic era (see Fig. 32-24). These early mammals remained a minor component of life on Earth for almost 150 million years and then rapidly diversified during the Cenozoic era (from 66 mya to the present). Fossil evidence indicates that the first primates with traits characteristic of modern primates appeared by the Early Eocene epoch about 56 mya. These early primates had digits with nails, and their eyes were directed somewhat forward on the head. Several novel adaptations evolved in early primates that allowed them to live in trees. One of the most significant primate features is that each limb has five highly flexible digits: four lateral digits (fingers) plus a partially or fully opposable first digit (thumb and, in many primates, big toe; FIG. 22-1). An opposable thumb positions the fingers opposite the thumb, enabling primates to grasp objects such as tree branches with precision. Nails (instead of claws) provide a protective covering for the tips of the digits, and the fleshy pads at the ends of the digits are sensitive to touch. Another arboreal feature is long, slender limbs that rotate freely at the hips and shoulders, giving primates full mobility to climb and search for food in the treetops. Having eyes located in the front of the head lets primates integrate visual information from both eyes simultaneously;

KEY POINT

Hand

Primates have five grasping digits at the end of each limb, and the thumb or first toe is often partially or fully opposable.

Foot

(a) Lemur (Eulemur mongoz) Hand

Foot

(c) Woolly spider monkey (Brachyteles arachnoides)

Hand

Foot

(b) Tarsier (Tarsius spectrum) Hand

Foot

(d) Gorilla (Gorilla gorilla)

Adapted from Schultz, A. H. The Life of Primates. Weidenfeld & Nicholson, London, 1969.

FIGURE 22-1 Right hands and feet of selected primates Figures not drawn to scale.

22.2 PRIMATE CLASSIFICATION ■ ■ LEARNING OBJECTIVES 2 3

List the three suborders of primates and give representative examples of each. Distinguish among anthropoids, hominoids, and hominins.

Now that we have surveyed the general characteristics of primates, let us look at how they are classified. Many biologists currently divide the order Primates into three groups, or suborders (FIG. 22-2). The suborder Prosimii includes lemurs, galagos, and lorises; the suborder Tarsiiformes includes tarsiers; and the suborder Anthropoidea includes anthropoids (monkeys, apes, and humans). All lemurs are restricted to the island of Madagascar off the coast of Africa. Because of extensive habitat destruction and hunt-

KEY POINT Suborder Prosimii

Lemurs

ing, they are very endangered. Lorises, which live in tropical areas of Southeast Asia and Africa, resemble lemurs in many respects, as do galagos, which live in sub-Saharan Africa. Lemurs, lorises, and galagos have retained several early mammalian features, such as elongated, pointed faces. Tarsiers are found in rain forests of Indonesia and the Philippines (FIG. 22-3). They are small primates (about the size of a small rat) and are very adept leapers. These nocturnal primates resemble anthropoids in several ways, including their shortened snouts and forward-pointing eyes.

Suborder Anthropoidea includes monkeys, apes, and humans Anthropoid primates arose during the Middle Eocene epoch, at least 45 mya. Several different fossil anthropoids have been

Several kinds of data support the hypothesis that chimpanzees are the closest living relative of humans.

Suborder Tarsiiformes

Tarsiers

Suborder Anthropoidea (Anthropoids) Hominoids Lesser Apes Great Apes

New World monkeys

Old World monkeys

Gibbons

Orangutans

Gorillas

Common anthropoid ancestor

Chimpanzees

Humans

Common hominoid ancestor

Common primate ancestor

FIGURE 22-2 Animated Primate evolution This branching diagram, called a cladogram, shows evolutionary relationships among living primates, based on current scientific evidence. The nodes (circles) represent branch points where a species splits into two or more lineages. The divergence of orangutans from the ape/

hominin line occurred some 12 mya to 16 mya. Gorillas separated from the chimpanzee/hominin line an estimated 8 mya, and the hominin (human) lineage diverged from that of chimpanzees about 6 mya. (Figures not drawn to scale.)

C. C. Lockwood/DRK Photo

identified, from Asia and North Africa, and there is a growing consensus about the relationships of these fossil groups to one another and to living anthropoids. Evidence indicates that anthropoids originated in Africa or Asia. The oldest known anthropoid fossils, such as 42-million-year-old Eosimias, have been found in China and Myanmar. Given details about their dentition and the few bones that have been discovered, scientists infer that Eosimias and other ancestral anthropoids were small, insect-eating arboreal primates that were active during the day. Once they evolved, anthropoids quickly spread throughout Europe, Asia, and Africa and arrived in South America much later. (Paleoanthropologists date the oldest known South American primate, Branisella, from Bolivia, at 26 million years old.) One significant difference between anthropoids and other primates is the size of their brains. The cerebrum, in

(a) New World monkey. The white-faced monkey (Cebus capucinus) has a prehensile tail and a flattened nose with nostrils directed to the side.

FIGURE 22-4 New World and Old World monkeys

Tierbild Okapia/Photo Researchers, Inc.

Frans Lanting/Minden Pictures

FIGURE 22-3 A tarsier The huge eyes of the tarsier (Tarsius bancanus) help it find insects, lizards, and other prey when it hunts at night. When a tarsier sees an insect, it pounces on it and grasps the prey with its hands. Tarsiers live in the rain forests of Indonesia and the Philippines.

particular, is more developed in monkeys, apes, and humans, where it functions as a highly complex center for learning, voluntary movement, and interpretation of sensation. Monkeys are generally diurnal (active during the day) tree dwellers. They tend to eat fruit and leaves, with nuts, seeds, buds, insects, spiders, birds’ eggs, and even small vertebrates playing a smaller part in their diets. The two main groups of monkeys, New World monkeys and Old World monkeys, are named for the hemispheres where they diversified. Monkeys in South and Central America are called New World monkeys, whereas monkeys in Africa, Asia, and Europe are called Old World monkeys. New and Old World monkeys have been evolving separately for tens of millions of years. One of the most important and interesting questions in anthropoid evolution concerns how monkeys arrived in South America. Africa and South America had already drifted apart. (Continental drift is presented in Figure 18-11.) The ancestors of New World monkeys may have rafted from Africa to South America on floating masses of vegetation. The South Atlantic Ocean would have been about half as wide as it is today, and islands could have provided “stepping stones.” Once established in the New World, these monkeys rapidly diversified. New World monkeys are restricted to Central and South America and include marmosets, capuchins, howler monkeys, squirrel monkeys, and spider monkeys. New World monkeys are arboreal, and some have long, slender limbs that permit easy movement in the trees (FIG. 22-4a). A few have prehensile tails capable of wrapping around branches and serving as fifth limbs. Some New World monkeys have shorter thumbs, and in certain cases the thumbs are totally absent. Their facial anatomy differs from that of the Old

(b) Old World monkey. The Anubis baboon (Papio anubis) is native to Africa. Note that its nostrils are directed downward.

World monkeys; New World monkeys have flattened noses with the nostrils opening to the side. They live in groups and engage in complex social behaviors. Old World monkeys are distributed in tropical parts of Africa and Asia. In addition to baboons and macaques (pronounced “muh-kacks′ ”), the Old World group includes guenons, mangabeys, langurs, and colobus monkeys. Most Old World monkeys are arboreal, although some, such as baboons and macaques, spend much of their time on the ground (FIG. 22-4b). The ground dwellers, which are quadrupedal (four footed; they walk on all fours), arose from arboreal monkeys. None of the Old World monkeys has a prehensile tail, and some have extremely short tails. They have a fully opposable thumb, and unlike the New World monkeys, their nostrils are closer together and directed downward. Old World monkeys are intensely social animals.

Apes are our closest living relatives Old World monkeys shared a common ancestor with the hominoids, a group composed of apes and hominins (humans and their ancestors; also called hominids). A fairly primitive anthropoid, discovered in Egypt, was named Aegyptopithecus (FIG. 22-5a). A cat-sized, forest-dwelling arboreal monkey with a few apelike characteristics, Aegyptopithecus lived during the Oligocene epoch, approximately 34 mya. Fossil evidence indicates that apes and Old World monkeys had diverged between 25 mya and 23 mya, although a 2004 genetic

analysis of living apes and monkeys suggests that this divergence may have occurred even earlier. During the Miocene epoch, as many as 100 species of apes lived in Africa, Asia, and Europe. Paleoanthropologists have discovered the oldest fossils with hominoid features in East Africa, mostly in Kenya. Proconsul, for example, appeared early in the Miocene epoch, about 20 mya (FIG. 22-5b). It had a larger brain than that of monkeys, apelike teeth and diet (fruits), but a monkeylike body. At least 30 other early hominoid species lived during the Miocene epoch, but most of them became extinct and were not the common ancestor of modern apes and humans. Miocene fossils of forest-dwelling, chimpanzee-sized apes called dryopithecines, which lived about 15 mya, are of special interest because this hominoid lineage may have given rise to modern apes as well as to humans (FIG. 22-5c). The dryopithecines, such as Dryopithecus, Kenyapithecus, and Morotopithecus, were distributed widely across Europe, Africa, and Asia. As the climate gradually cooled and became drier, their range became more limited. These apes had highly modified bodies for swinging through the branches of trees, although there is also evidence that some of them may have left the treetops for the ground as dense forest gradually changed into open woodland. Many questions about the relationships among the various early apes have been generated by the discovery of these and other Miocene hominoids. As future fossil finds are evaluated, they may lead to a rearrangement of ancestors in the hominoid family tree. Five genera of hominoids exist today: gibbons (Hylobates), orangutans (Pongo), gorillas (Gorilla), chimpanzees (Pan), and humans (Homo). Gibbons are known informally as lesser apes; orangutans, gorillas, and chimpanzees are known as great apes. Molecular evidence indicates a close relationship between humans and the great apes, particularly chimpanzees.

(a) Fossils of Aegyptopithecus, a fairly primitive anthropoid, were discovered in Egypt.

(b) Skeletal reconstruction of Proconsul. (The reconstructed parts are white.) This anthropoid had the limbs and body proportions of a monkey but lacked a tail, like all apes. (c) Dryopithecus, a more advanced ape, may have been ancestral to modern hominoids.

FIGURE 22-5 Ape evolution (Figures not drawn to scale.)

Review ■ ■

BIOS/Peter Arnold, Inc.

(b) An orangutan (Pongo pygmaeus) mother and baby. Orangutan anatomy is adapted to living in trees.

(a) A mother white-handed gibbon (Hylobates lar ) nurses her baby. Gibbons are extremely acrobatic and often move through the trees by brachiation.

(c) A young lowland gorilla (Gorilla gorilla) in knuckle-walking stance. Gorillas spend most of the day eating plants.

FIGURE 22-6 Apes

What are the three suborders of primates? How can you distinguish between anthropoids and hominoids?

Michael K. Nichols/NGS/Getty Images

Nancy Adams/Tom Stack & Associates

Molecular data provide clues about evolution of anthropoid primates Evidence of the close relatedness of orangutans, gorillas, chimpanzees, and humans is abundant at the molecular level. The amino acid sequence of the chimpanzee’s hemoglobin is identical to that of the human; hemoglobin molecules of the gorilla and rhesus monkey differ from the human’s by 2 and 15 amino acids, respectively. DNA sequence analyses indicate that chimpanzees are likely to be our nearest living relatives among the apes. (Figure 18-18 shows the percent divergence in a noncoding sequence on the b-globin gene between various primates, including the apes, and humans.) Researchers plan to sequence the genomes of many primates, including all of the apes. The chimpanzee genome was sequenced in 2005, and the macaque (an Old World monkey) in 2007. Macaques are thought to have split from the anthropoid lineage about 25 mya, so comparing their genome to those of chimpanzees and humans helps researchers identify the genetic changes that evolved only in the anthropoid lineage. Molecular evidence suggests that orangutans may have diverged from the gorilla, chimpanzee, and hominin lines about 14 mya. Gorillas may have diverged from the chimpanzee and hominin lines some 8 mya, whereas chimpanzee and hominin lines probably separated about 6 mya to 7 mya.

Joe McDonald/Visuals Unlimited, Inc.

Gibbons are natural acrobats that can brachiate, or arm-swing, with their weight supported by one arm at a time (FIG. 22-6a). Orangutans are also tree dwellers, but chimpanzees and especially gorillas have adapted to life on the ground (FIG. 22-6b–d). Gorillas and chimpanzees have retained long arms typical of brachiating primates but use these to assist in quadrupedal walking, sometimes known as knuckle walking because of the way they fold (flex) their digits when moving. Like humans, apes lack tails, a characteristic that makes them easy to distinguish from monkeys. They are generally much larger than monkeys, although gibbons are a notable exception.

(d) A mother bonobo chimpanzee (Pan paniscus ) holds her sleeping baby. Bonobos are endemic to a single country, the Democratic Republic of Congo.

22.3 HOMININ EVOLUTION ■ ■ LEARNING OBJECTIVES 4 Describe skeletal and skull differences between apes and hominins. 5 Briefly describe the following early hominins: Orrorin, Ardipithecus, and Australopithecus anamensis, A. afarensis, and A. africanus.

6 Distinguish among the following members of genus Homo: H. habilis,

7

H. ergaster, H. erectus, H. antecessor, H. heidelbergensis, H. neanderthalensis, and H. sapiens. Discuss the origin of modern humans.

Scientists have a growing storehouse of hundreds of hominin fossils, which provide useful data about general trends in the body design, appearance, and behavior of ancestral humans. For example, before their brains enlarged, early hominins clearly adopted a bipedal (two-footed) posture. Despite the wealth of fossil evidence, scientists continue to vigorously debate interpretations of hominin characteristics, classification, and evolution, and new discov-

Simply curved spine

eries raise new questions. As in other scientific fields, ideas about hominin evolution are influenced by the different perspectives of the various workers studying it. The lack of scientific consensus regarding certain aspects of hominin evolution is, therefore, an expected part of the scientific process. Evolutionary changes from the earliest hominins to modern humans are evident in some of the characteristics of the skeleton and skull. Compared with the ape skeleton, the human skeleton shows distinct differences that reflect humans’ ability to stand erect and walk on two feet (FIG. 22-7). These differences also reflect the habitat change for early hominins, from an arboreal existence in the forest to a life spent at least partly on the ground. The curvature of the human spine provides better balance and weight distribution for bipedal locomotion. The human pelvis is shorter and broader than the ape pelvis, allowing better attachment of muscles used for upright walking. (The shape of the human pelvis also allows the birth of offspring with large brains.) In apes the foramen magnum, the hole in the base of the skull for the spinal cord, is located in the rear of the skull. In contrast, the

Foramen magnum at the center base of skull

Complex curvature of human spine

Foramen magnum at the center rear of skull

Tall, narrow pelvis (front view)

Shorter, broader pelvis (front view)

First toe not opposable, and all toes aligned

First toe not aligned with others Gorilla skeleton

FIGURE 22-7 Gorilla and human skeletons When gorilla and human skeletons are compared, the skeletal adaptations for bipedalism in humans become apparent.

Human skeleton

human foramen magnum is centered in the skull base, positioning the head for erect walking. An increase in the length of the legs relative to the arms, and alignment of the first toe with the rest of the toes, further adapted the early hominins for bipedalism. Another major trend in hominin evolution was an increase in brain size relative to body size (FIG. 22-8). The ape skull has prominent supraorbital ridges above the eye sockets, whereas modern human skulls lack these ridges. Human faces are flatter than those of apes, and the jaws are different. The arrangement of teeth in the ape jaw is somewhat rectangular, compared with a rounded, or U-shaped, arrangement in humans. Apes have larger front teeth (canines and incisors) than do humans, and their canines are especially large. Gorillas and orangutans also have larger back teeth (premolars and molars) than do humans. We now examine some of the increasing number of fossil hominins in the human lineage. As you read the following descriptions of human evolution, keep in mind that much of what is discussed is open to reinterpretation and major revision as additional discoveries are made. Although we present human evolution in a somewhat linear fashion, from ancient hominins to anatomically and behaviorally modern humans, the human family tree is not a single trunk

Supraorbital ridge

Incisors Rectangular shape

(a) The ape skull has a pronounced supraorbital ridge.

Incisors U shape

(b) The human skull is flatter in the front and has a pronounced chin. The human brain, particularly the cerebrum (purple), is larger than that of an ape, and the human jaw is structured so that the teeth are arranged in a U shape. Human canines and incisors are also smaller than those of apes.

FIGURE 22-8 Gorilla and human heads

but has several branches (FIG. 22-9). Perhaps it is most useful to think of the known hominin fossils as a sampling of human evolution instead of a continuous sequence. Homo sapiens is the only species of hominin in existence today, but more than one hominin species coexisted at any given time for most of the past 4 million years. In addition, do not make the mistake of thinking that your smaller-brained ancestors were inferior to yourself. Ancestral hominins were evolutionarily successful in that they were well adapted to their environment and survived for millions of years.

The earliest hominins may have lived 6 mya to 7 mya Hominin evolution began in Africa. Although most hominin fossils have been discovered in Ethiopia and Kenya, in 2002 French paleontologist Michel Brunet and an international team made a stunning discovery in a dry lake bed in Chad, which is in central Africa. The fossil, which has been reliably dated at 6 million to 7 million years old, may be the earliest known hominin. Viewed from the back, the skull of Sahelanthropus, with its small braincase, resembles that of a chimpanzee. However, viewed from the front, the face and teeth have many characteristics of larger-brained human ancestors. Most paleoanthropologists place Sahelanthropus close to the base of the human family tree—that is, close to when genetic analyses suggest the last common ancestor of hominins and chimpanzees existed. The discovery of Sahelanthropus is important in its own right, but it is also significant because it shows that early hominins exhibited more variation and lived over a larger area of Africa than had been originally hypothesized. Orrorin, discovered in 2000 in Kenya, may also represent one of the earliest known hominins. The sediments in which fossils of Orrorin were found have been reliably dated at 6 million years old. In 2008, researchers studying the fossil leg bones of Orrorin concluded that it walked upright and was bipedal. As with many aspects of human evolution, scientists have differing interpretations of Sahelanthropus and Orrorin. For example, some paleoanthropologists hypothesize that Sahelanthropus was a forerunner of modern apes, specifically the gorilla, and not one of the earliest humans. Other researchers think the similar features of Sahelanthropus, Orrorin, and Ardipithecus (discussed in the next section) fossils mean that they are all members of the same genus, Ardipithecus. This point remains controversial, largely because there are currently no skeletal bones (leg, pelvis, and foot bones) to indicate if Sahelanthropus walked upright, a hallmark characteristic of hominins. (A 2005 reconstruction of the skull has features that strongly suggest Sahelanthropus was bipedal.) Future fossil discoveries and additional analyses of existing fossils should help clarify the evolutionary relationships among Sahelanthropus, Orrorin, and Ardipithecus.

Ardipithecus, Australopithecus, and Paranthropus are australopithecines, or “southern man apes” After Sahelanthropus and Orrorin, the next-oldest hominin belongs to the genus Ardipithecus, which lived in eastern Africa between

KEY POINT

Paleoanthropologists do not completely agree about certain specific details of the human family tree and hold many possible interpretations of classification lines of descent.

Homo neanderthalensis

Orrorin

Australopithecus anamensis

Ardipithecus ramidus

Ardipithecus kadabba

Australopithecus africanus

Homo heidelbergensis

Homo ergaster

Australopithecus afarensis

Homo antecessor

Homo sapiens

Homo habilis Homo erectus Paranthropus robustus

Key:

Paranthropus aethiopicus

Duration of species Line of descent is well-supported Less-certain line of descent

6

5

Paranthropus boisei

4 3 Millions of years ago (mya)

2

1

Present

FIGURE 22-9 One interpretation of human evolution This interpretation, compiled by the author from multiple sources, will likely change as new evidence comes to light.

4 mya and 6 mya. Although not as primitive as Sahelanthropus, Ardipithecus is close to the “root” of the human family tree. The shape of the toe bones suggests that Ardipithecus walked upright. Two species of Ardipithecus have been described. The canine teeth of the earliest species, A. kadabba, have certain primitive features similar to those of Sahelanthropus and Orrorin. The specific epithet kadabba comes from Afar words meaning “basal family ancestor.” Ardipithecus ramidus appeared about 4.8 mya. A remarkably complete fossil of a female A. ramidus, named Ardi, was scientifically described in 2009. Ardipithecus gave rise to Australopithecus, a genus that includes several species that lived between 4 mya and 1 mya. Both Ardipithecus and Australopithecus had longer arms, shorter legs, and smaller brains relative to those of modern humans. Hominins that existed between 4.2 mya and 3.9 mya are assigned to the species Australopithecus anamensis, first named by paleoanthropologist Meave Leakey and her co-workers in 1995 from fossils discovered in eastern Africa. This hominin species, which has a mixture of apelike and humanlike features, presumably evolved from Ardipithecus ramidus. A comparison of male and female Australopithecus anamensis body sizes and canine teeth reveals sexual dimorphism, marked phenotypic differences between the two sexes of the same species. (The modern-day gorilla is sexually dimorphic.) The back teeth and jaws of A. anamensis are

larger than those of modern chimpanzees, whereas the front teeth are smaller and more like those of later hominins. A fossil leg bone, the tibia, indicates that A. anamensis had an upright posture and was bipedal, although it also may have foraged in the trees. Thus, bipedalism occurred early in human evolution and appears to have been the first human adaptation. Australopithecus afarensis, another primitive hominin, probably evolved directly from A. anamensis. Many fossils of A. afarensis skeletal remains have been discovered in Africa, including a remarkably complete 3.2-million-year-old skeleton, nicknamed Lucy, found in Ethiopia in 1974 by a team led by U.S. paleoanthropologist Donald Johanson (FIG. 22-10). In 1978, British paleoanthropologist Mary Leakey and co-workers discovered beautifully preserved fossil footprints of three A. afarensis individuals who walked more than 3.6 mya. In 1994, paleoanthropologists found the first adult skull of A. afarensis. The skull, characterized by a relatively small brain, pronounced supraorbital ridges, a jutting jaw, and large canine teeth, is an estimated 3 million years old. It is probable that A. afarensis did not construct tools or make fires because no evidence of tools or fire has been found at fossil sites. Many paleoanthropologists think that A. afarensis gave rise to several australopithecine species, including Australopithecus africanus, which may have appeared as early as 3 mya. The first A. africanus fossil was discovered in South Africa in 1924, and since

© Lionel Bret/Photo Researchers, Inc.

Homo habilis is the oldest member of genus Homo

FIGURE 22-10 Australopithecus afarensis This artist’s rendition of a female australopithecine and child is based on the fossils of Lucy, discovered in 1974.

then hundreds have been found. This hominin walked erect and possessed hands and teeth that were distinctly humanlike. Given the characteristics of the teeth, paleoanthropologists think that A. africanus ate both plants and animals. Like A. afarensis, it had a small brain, more like that of its primate ancestors than of presentday humans. Three australopithecine species (Paranthropus robustus from South Africa, and P. aethiopicus and P. boisei, both from East Africa) were larger than A. africanus and had extremely large molars, very powerful jaws, relatively small brains, and bony skull crests. Most females lacked the skull crests and had substantially smaller jaws, another example of sexual dimorphism in early hominins. The teeth and jaws suggest a diet, perhaps of tough roots, nuts, and seeds, that would require powerful grinding. (However, a 2008 study of fossil teeth of P. boisei found that the wear on the teeth suggested a softer diet, such as fruits. P. boisei apparently ate soft food during times of plenty but could eat hard items during food shortages.) These so-called robust australopithecines may or may not be closely related but are generally thought to represent evolutionary offshoots, or side branches, of human evolution. The first robust australopithecine, P. aethiopicus, appeared about 2.5 mya. The actual number of australopithecine species for which fossil evidence has been found is under debate. In some cases, differences in the relatively few skeletal fragments could indicate either variation among individuals within a species or evidence of separate species.

The first hominin to have enough uniquely human features to be placed in the same genus as modern humans is Homo habilis. It was first discovered in the early 1960s at Olduvai Gorge in Tanzania. Since then, paleoanthropologists have discovered other fossils of H. habilis in eastern and southern Africa. Homo habilis was a small hominin with a larger brain and smaller premolars and molars than those of the australopithecines. This hominin appeared approximately 2.5 mya and persisted for about 0.9 million years. Fossils of H. habilis have been found in numerous areas in Africa. These sites contain primitive tools, stones that had been chipped, cracked, or hammered to make sharp edges for cutting or scraping. Oldowan pebble choppers and flakes, for example, were probably used to cut through animal hides to obtain meat and to break bones for their nutritious marrow. The relationship between the australopithecines and H. habilis is not clear. Using physical characteristics of their fossilized skeletons as evidence, many paleoanthropologists have inferred that the australopithecines were ancestors of H. habilis. Some researchers do not think that H. habilis belongs in the genus Homo, and they suggest it should be reclassified as Australopithecus habilis. Discoveries of additional fossils may help clarify these relationships.

Homo ergaster may have arisen from H. habilis Initially, fossils that are now classified as Homo ergaster were considered by some scientists as early H. erectus (discussed in next section). With the discovery of additional fossils, however, many scientists hypothesized that the fossils classified as H. erectus really represent two species, Homo ergaster, an earlier African species, and H. erectus, a later eastern Asian offshoot. The best-known fossils of H. ergaster, which existed from 2.0 to 1.4 mya, come from the Lake Turkana region in Kenya. Researchers who support the split speculate that H. ergaster may be the direct ancestor of later humans, whereas H. erectus may be an evolutionary dead end.

Homo erectus probably evolved from H. ergaster Investigators found the first fossil evidence of Homo erectus in Indonesia in the 1890s. Since then, searchers have found numerous fossils of H. erectus throughout Africa and Asia. Paleoanthropologists think that H. erectus originated in Africa about 1.7 mya and then spread to Europe and Asia. Peking man and Java man, discovered in Asia, were later examples of H. erectus, which existed until at least 200,000 years ago; some populations of H. erectus may have persisted more recently. Homo erectus was taller than H. habilis. Its brain, which was larger than that of H. habilis, got progressively larger during the course of its evolution. Its skull, although larger, did not possess totally modern features, retaining the heavy supraorbital ridge and projecting face that are more characteristic of its ape ancestors (FIG. 22-11). Homo erectus is the first hominin to have fewer differences between the sexes. The increased intelligence associated with a larger brain enabled these early humans to make more advanced stone tools, known as Acheulean tools, including hand axes and other imple-

© Javier Trueba/Madrid Scientific Films/Photo Researchers, Inc.

Receding forehead

Ken Mowbray, AMNH

Pronounced supraorbital ridge

Projecting face/jaws

FIGURE 22-11 Homo erectus skull from China The reconstructed parts are white. Note the receding forehead, pronounced supraorbital ridge, and projecting face and jaws.

ments that scientists have interpreted as choppers, borers, and scrapers (FIG. 22-12). Their intelligence also allowed these humans to survive in cold areas. Homo erectus obtained food by hunting or scavenging and may have worn clothing, built fires, and lived in caves or shelters. Evidence of weapons (spears) has been unearthed at H. erectus sites in Europe. (For the remarkable story of an evolutionary offshoot from H. erectus, see Inquiring About: The Smallest Humans.)

Archaic humans date from about 1.2 mya to 200,000 years ago Archaic humans are regionally diverse descendants of H. ergaster that lived in Africa, Asia, and Europe from about 1.2 mya to 200,000 years ago. The brains of archaic humans were essentially the same size as our brains, although their skulls retained some ancestral characters. Populations of archaic humans had rich and varied cultures (learned traditions) that included making many kinds of tools and objects with symbolic and ceremonial meaning. Some researchers classify the oldest archaic human fossils discovered in Europe (in Spain) as Homo antecessor. Homo antecessor existed from about 1.2 mya to 800,000 years ago. Sites where H. antecessor fossils have been found show numerous cuts on human bones that suggest these early humans practiced cannibalism. Homo heidelbergensis, which appeared about 600,000 years ago and existed until about 300,000 years ago, may have descended

FIGURE 22-12 Acheulean tool This stone tool is a typical Acheulean tool. It was made about 800,000 years ago by striking both sides of the biface (shown) to make a teardrop shape. The tool was found with other fossils and tools in Gran Dolina, Sierra de Atapuerca, Spain.

from H. antecessor. Fossils of H. heidelbergensis were first discovered in Germany about 100 years ago. This extinct hominin had a larger brain—similar in size to that of modern humans—than H. ergaster or H. antecessor. Some scientists do not recognize H. antecessor and suggest H. antecessor fossils should be classified as H. heidelbergensis. Classification of H. heidelbergensis is also controversial because many (but not all) of the scientists studying its fossil remains think that both Neandertals and modern humans descended from it.

Neandertals appeared approximately 250,000 years ago Fossils of Neandertals were first discovered in the Neander Valley in Germany.1 They lived throughout Europe and western Asia from about 250,000 to 28,000 years ago. Although at one time many scientists considered Neandertals as a subspecies of H. sapiens, increasing evidence suggests Neandertals were a separate species from modern humans, Homo neanderthalensis.

1

Neandertal was formerly spelled Neanderthal. The silent h has been dropped in modern German but not in the scientific name.

In qu i ri n g A b o u t

Evolutionarily speaking, what are “hobbits”? In 2004, paleoanthropologists reported a startling discovery—fossils of seven humans in a cave on the Indonesian island of Flores. The finding was completely unexpected because the fossil bones and teeth were from adult humans that were about 1 m (about 3 ft) tall. The tiny humans, which have been dubbed “hobbits” by the media, may represent a new species, Homo floresiensis. Although these hominins had small, ape-sized brains, sophisticated stone tools, fireplaces, and charred bones found with the fossils indicate that H. floresiensis was capable of complex thinking and activities. The charred bones were primarily from pygmy elephants (Stegodon) and Flores giant rats (Papagomys) and provide clues about the food that H. floresiensis hunted and killed (see figure). Based on several dating methods, H. floresiensis is thought to have existed from about 38,000 years ago to as recently as 12,000 years ago, when it went extinct. Detailed studies of the H. floresiensis braincase, published in 2005, revealed that it was similar in many ways to the larger brain of H. erectus, which is known to have lived on nearby islands. Many researchers have concluded from this and other evi-

dence that H. floresiensis was an evolutionary offshoot of H. erectus. Why was H. floresiensis so small? Biologists have often observed two evolutionary trends of mammals living on remote islands: large mammals tend to evolve into much smaller species, and small mammals tend to evolve into much larger species. It is not unreasonable to assume that the small population of H. erectus ancestors that colonized Flores, perhaps by rafting to the island on a log, underwent evolutionary dwarfing over the thousands of years they were isolated on Flores. (If H. erectus had been a frequent visitor to Flores, gene flow between the indigenous population and the visitors would have prevented the indigenous population from becoming measurably smaller.) Not everyone agrees with the hypothesis that H. floresiensis is a separate species of tiny humans. Some researchers think the fossils have not been identified properly. In a study published in 2006, these researchers suggest that the fossils are from H. sapiens individuals suffering from a rare genetic defect (microcephaly) that causes small brains and bodies. The research team that unearthed the fossils has returned to the cave and will continue excavations. It is anticipated

These early humans had short, sturdy builds. Their faces projected slightly, their chins and foreheads receded, they had heavy supraorbital ridges and jawbones, and their brains and front teeth were larger than those of modern humans. They had large nasal cavities and receding cheekbones. Scientists have suggested that the large noses provided larger surface areas in Neandertal sinuses, enabling them to better warm the cold air of Ice Age Eurasia as inhaled air traveled through the head to the lungs. Neandertal tools, known as Mousterian tools, were more sophisticated than those of H. erectus (FIG. 22-13). Studies of Neandertal sites indicate that Neandertals hunted large animals. The existence of skeletons of elderly Neandertals and of some with healed fractures may demonstrate that Neandertals cared for the aged and the sick, an indication of advanced social cooperation. They apparently had rituals, possibly of religious significance, and sometimes buried their dead. The disappearance of the Neandertals about 28,000 years ago is a mystery that has sparked debate among paleoanthropologists. Other groups of H. sapiens with more modern features coexisted with the Neandertals for several thousand years. Perhaps the other humans outcompeted or exterminated the Neandertals, leading to their extinction. It is also possible that the Neandertals interbred with these humans, diluting their features beyond recognition. Analysis of mitochondrial DNA (mtDNA) contributes useful data for such controversies. Each of the several hundred mitochon-

Peter Schouten/National Geographic Society/University of Wollongong

THE SMALLEST HUMANS

that future discoveries will help scientists answer the many questions we have about Earth’s smallest humans. Meanwhile, the debate about the origin of H. floresiensis continues.

dria within a cell has about 10 copies of a small loop of DNA that codes for transfer RNAs, ribosomal RNAs, and certain respiratory enzymes. The mtDNA is transmitted only through the maternal line because eggs, not sperm, contribute mitochondria. Because mtDNA mutates more rapidly than nuclear DNA, mtDNA is a sensitive indicator of evolution. Researchers have extracted and evaluated the mtDNA from seven separate Neandertals. Its sequence differs significantly from all modern human mtDNA sequences, although it is more similar to human than to chimpanzee mtDNA. This finding suggests that Neandertals are an evolutionary dead end that did not interbreed appreciably with more modern humans. However, the amount of interbreeding that may have occurred between Neandertals and anatomically modern humans remains controversial.

Scientists have reached a near consensus on the origin of modern Homo sapiens Homo sapiens with anatomically modern features existed in Africa about 195,000 years ago. As these anatomically modern humans dispersed around the world, they displaced other hominins such as H. erectus and H. neanderthalensis. By about 30,000 years ago, anatomically modern humans were the only members of genus Homo remaining, excluding small, isolated populations. Both recent fossil finds and extensive molecular analyses have promoted the (recent) out-of-Africa model (also called Recent

© The Natural History Museum, London

Africa Origin, or RAO) as the main explanation for the origin of modern humans living around the world. New data include the discovery of the earliest fossilized remains of modern H. sapiens in Africa and analyses of DNA from mitochondria, the Y chromosome, and autosomes. In 2005, fossils of the earliest known modern H. sapiens were reliably dated at 195,000 years old; these fossils were found in southern Ethiopia. No H. sapiens fossils of comparable age have been found in Europe or Asia. The earliest fossils of anatomically modern H. sapiens in Europe and most parts of Asia date to 45,000 to 40,000 years ago. One exception is in Israel, where H. sapiens fossils have been dated at 100,000 years before present. Molecular anthropology, the comparison of biological molecules from present-day individuals of regional human populations, provides clues that help scientists unravel the origin of modern humans and trace human migrations. A series of recent genetic studies of mtDNA, Y chromosome, and autosomal DNA has strengthened the case for Africa as the birthplace of modern humans (FIG. 22-14). These studies provide detailed comparisons of DNA in present-day populations around the world as well as ancient DNA extracted from Neandertal and early H. sapiens remains.

Review FIGURE 22-13 Mousterian tool

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Mousterian tools are named after a Neandertal site in Le Moustier, France. Mousterian tools include a variety of skillfully made stone tools; this tool, with its sharp edges, was probably used to scrape hides and/or to form the point of a spear. Although Neandertals had spears, they did not develop bows and arrows and so had to get close to their prey when hunting.

LLY22

M17 M170 M9

M172, M135 M304 M96 YAP M60 M2

■ ■ ■

How do the skulls of apes and humans differ? How does an ape skeleton differ from a human skeleton? How do australopithecines and Homo differ? How can you distinguish between H. habilis and H. erectus? How do H. neanderthalensis and H. sapiens differ?

M242

M173 M343

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M45

M201 M20

M174 M3

M122

M130 M89 M168 M69

M175 M4

Origin M91 M130

FIGURE 22-14 The path of human migrations, based on Y chromosome data Homo sapiens originated in eastern Africa and migrated from there throughout the world. (Source: Stix, G. "Traces of a Distant Past." Scientific American, p. 59, July 2008. Author credits information in National Geographic maps.)

22.4 CULTURAL CHANGE ■ ■ LEARNING OBJECTIVE 8

Describe the impact of human culture on the biosphere.

Genetically speaking, humans are not very different from other primates. At the level of our DNA sequences, we are roughly 98% identical to gorillas and 99% identical to chimpanzees. Our relatively few genetic differences, however, give rise to several important distinguishing features, such as greater intelligence and the ability to capitalize on it by transmitting knowledge from one generation to the next. Human culture is not inherited in the biological sense but instead is learned, largely through the medium of language.2 Human culture is dynamic and is modified as people obtain new knowledge. Human culture is generally divided into three stages: (1) the development of hunter–gatherer societies, (2) the development of agriculture, and (3) the Industrial Revolution. Early humans were hunters and gatherers who relied on what was available in their immediate environment. They were nomadic, and as the resources in a given area were exhausted or as the population increased, they migrated to a different area. These societies required a division of labor and the ability to make tools and weapons, which were needed to kill game, scrape hides, dig up roots and tubers, and cook food. Although scientists are not certain when hunting was incorporated into human society, they do know it declined in importance approximately 15,000 years ago. This may have been due to a decrease in the abundance of large animals, triggered in part by overhunting. A few isolated groups of hunter–gatherer societies, such as the Inuit of the northern polar region and the Mbuti of Africa, survived into the 21st century.

Development of agriculture resulted in a more dependable food supply Evidence that humans had begun to cultivate crops approximately 10,000 years ago includes the presence of agricultural tools and plant material at archaeological sites. Agriculture resulted in a more dependable food supply. Archaeological evidence suggests that agriculture arose in several steps. Although there is variation from one site to another, plant cultivation, in combination with hunting, usually occurred first. Animal domestication generally followed later, although in some areas, such as Australia, early humans did not domesticate animals. Agriculture, in turn, often led to more permanent dwellings because considerable time was invested in growing crops in one area. Villages and cities often grew up around the farmlands, but the connection between agriculture and the establishment of villages and towns is complicated by certain discoveries. For example, Abu

Hureyra in Syria was a village founded before agriculture arose. The villagers subsisted on the rich plant life of the area and the migrating herds of gazelle. Once people turned to agriculture, however, they seldom went back to hunting and gathering to obtain food. Other advances in agriculture include the domestication of animals, which people kept to supply food, milk, and hides. Archaeological evidence indicates that wild goats and sheep were probably the first animals to be domesticated in southwestern Turkey, northern Iraq, and Iran. In the Old World, people also used animals to prepare fields for planting. Another major advance in agriculture was irrigation, which began more than 5000 years ago in Egypt. Producing food agriculturally was more time-consuming than hunting and gathering, but it was also more productive. In hunter– gatherer societies, everyone shares the responsibility for obtaining food. In agricultural societies, fewer people are needed to provide food for everyone. Thus, agriculture freed some people to pursue other endeavors, including religion, art, and various crafts.

Human culture has had a profound impact on the biosphere Human culture has had far-reaching effects on both human society and on other organisms. The Industrial Revolution, which began in the 18th century, drew populations to concentrate in urban areas near centers of manufacturing. Advances in agriculture encouraged urbanization because fewer and fewer people were needed in rural areas to produce food for everyone. The spread of industrialization increased the demand for natural resources to supply the raw materials for industry. Human culture has permitted the human population, which reached 6.8 billion in 2009, to expand so dramatically that there are serious questions about Earth’s ability to support so many people indefinitely (see Chapter 53). According to the United Nations’ 2008 State of Food Insecurity, 923 million people lack access to the food needed to be healthy and lead productive lives. To further compound the problem, the UN projects that 2.6 billion additional people will be added to the world population by the year 2050. Human culture has resulted in large-scale disruption and degradation of the environment. Tropical rain forests and other natural environments are rapidly being eliminated. Soil, water, and air pollution occur in many places. Since World War II, soil degradation caused by poor agricultural practices, overgrazing, and deforestation has occurred in an area equal to 17% of Earth’s total vegetated surface area. Many species cannot adapt to the rapid environmental changes caused by humans and thus are becoming extinct. The decrease in biological diversity from species extinction is alarming (see Chapter 57). On a positive note, people are aware of the damage we are causing and have the intelligence to modify their behavior to improve these conditions. Education, including the study of biology, may help future generations develop greater environmental sensitivity.

2

Humans are not the only animals to have culture. Chimpanzees have primitive cultures that include tool-using techniques, hunting methods, and social behaviors that vary from one population to another. These cultural traditions are passed to the next generation by teaching and imitation; see Figures 1-18 and 52-13b.

Review ■

How has human culture affected planet Earth?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

22.1 (page 466) 1 Describe the structural adaptations that primates have for life in treetops. Primates are placental mammals that arose from small, arboreal (tree-dwelling), shrewlike mammals. Primates possess five grasping digits, including an opposable thumb or toe; long, slender limbs that move freely at the hips and shoulders; and eyes located in front of the head.

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22.2 (page 467) 2 List the three suborders of primates and give representative examples of each. Primates are divided into three suborders. The suborder Prosimii includes lemurs, galagos, and lorises. The suborder Tarsiiformes includes tarsiers. The suborder Anthropoidea includes anthropoids— monkeys, apes, and humans.

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Learn more about primate evolution by clicking on the figure in CengageNOW.

3 Distinguish among anthropoids, hominoids, and hominins. ■

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Anthropoids include monkeys, apes, and humans. Early anthropoids branched into two groups: New World monkeys and Old World monkeys. Hominoids include apes and humans. Hominoids arose from the Old World monkey lineage. There are four modern genera of apes: gibbons, orangutans, gorillas, and chimpanzees. The hominin line consists of humans and their ancestors.

22.3 (page 471) 4 Describe skeletal and skull differences between apes and hominins. ■

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Unlike ape skeletons, hominin skeletons have adaptations that reflect the ability to stand erect and walk on two feet. These adaptations include a complex curvature of the spine; a shorter, broader pelvis; repositioning of the foramen magnum to the base of the skull; and a first toe that is aligned with the other toes. The human skull lacks a pronounced supraorbital ridge, is flatter than ape skulls in the front, and has a pronounced chin. The human brain is larger than ape brains, and the jaw is structured so that the teeth are arranged in a U shape.

a hominin feature. The first Ardipithecus species, A. kadabba, appeared about 5.8 mya. Ardipithecus ramidus, which appeared about 4.8 mya, may have given rise to Australopithecus anamensis, which in turn probably gave rise to another primitive hominin, Australopithecus afarensis. Many paleoanthropologists think that A. afarensis gave rise to several australopithecine species, including Australopithecus africanus, Paranthropus spp., and possibly Homo habilis. 6 Distinguish among the following members of genus Homo: H. habilis, H. ergaster, H. erectus, H. antecessor, H. heidelbergensis, H. neanderthalensis, and H. sapiens. ■ Homo habilis was the earliest known hominin with some of the human features lacking in the australopithecines, including a slightly larger brain. Homo habilis fashioned crude tools from stone. ■ The best-known fossils of H. ergaster, which existed from 2.0 to 1.4 mya, come from Kenya. Homo ergaster may be the direct ancestor of later humans. ■ Homo erectus had a larger brain than H. habilis; made more sophisticated tools; and may have worn clothing, built fires, and lived in caves or shelters. Homo erectus, which is probably a later Asian offshoot of H. ergaster, appears to be an evolutionary dead end. ■ Archaic humans are regionally diverse descendants of H. ergaster that lived in Africa, Asia, and Europe from about 1.2 mya to 200,000 years ago. The brains of archaic humans were essentially the same size as our brains, although their skulls retained some ancestral characters, and they had rich and varied cultures. ■ Some researchers classify the oldest archaic human fossils discovered in Europe (in Spain) as Homo antecessor. Homo antecessor existed from about 1.2 mya to 800,000 years ago. ■ Homo heidelbergensis, which appeared about 600,000 years ago and existed until about 300,000 years ago, may have descended from H. antecessor. This extinct hominin had a larger brain—similar in size to that of modern humans—than either H. ergaster or H. antecessor. Many scientists studying H. heidelbergensis think that both Neandertals and modern humans descended from it. ■ Neandertals lived from about 250,000 to 30,000 years ago. Neandertals had short, sturdy builds; receding chins and foreheads; heavy supraorbital ridges and jawbones; larger front teeth; and nasal cavities with unusual triangular bony projections. Many scientists think that Neandertals were a separate species, Homo neanderthalensis. ■ Homo sapiens, anatomically modern humans, existed in Africa about 195,000 years ago. By about 30,000 years ago, anatomically modern humans were the only members of genus Homo remaining, excluding small, isolated populations. Learn more about Homo skulls by clicking on the figure in CengageNOW.

7 Discuss the origin of modern humans. ■

Learn more about monkey, gorilla, and human skeletons by clicking on the figure in CengageNOW.

5 Briefly describe the following early hominins: Orrorin, Ardipithecus, and Australopithecus anamensis, A. afarensis, and A. africanus. ■ Orrorin is an early hominin that arose about 6 mya. Researchers studying the fossil leg bones of Orrorin think that it walked upright and was bipedal (walked on two feet). Ardipithecus, Australopithecus, and Paranthropus species are often ■ referred to as australopithecines. Australopithecines were bipedal,

Recent fossil finds and extensive molecular analyses have promoted the (recent) out-of-Africa model as the main explanation for the origin of modern humans living around the world. New data include the discovery of the earliest fossilized remains of modern H. sapiens in Africa and analyses of DNA from mitochondria, the Y chromosome, and autosomes.

22.4 (page 478) 8 Describe the impact of human culture on the biosphere. ■

Large human brain size makes possible the transmission of knowledge from one generation to the next. Two significant advances in human culture were the development of agriculture and the Industrial Revolution.

T E S T YO U R U N D E R S TA N D I N G 1. The first primates evolved from (a) shrewlike monotremes (b) therapsids (c) shrewlike placental mammals (d) tarsiers (e) shrewlike marsupials 2. The anthropoids are more closely related to _____ than to _____. (a) tarsiers; lemurs (b) lemurs; monkeys (c) tree shrews; tarsiers (d) lemurs; tarsiers (e) tree shrews; monkeys 3. With what group do hominoids share the most recent common ancestor? (a) Old World monkeys (b) New World monkeys (c) tarsiers (d) lemurs (e) lorises and galagos 4. The _____ in humans is centered at the base of the skull, positioning the head for erect walking. (a) supraorbital ridge (b) foramen magnum (c) pelvis (d) bony skull crest (e) femur 5. Scientists collectively call humans and their immediate ancestors (a) mammals (b) primates (c) anthropoids (d) hominoids (e) hominins 6. The earliest hominin that scientists placed in the genus Homo is (a) H. habilis (b) H. ergaster (c) H. erectus (d) H. heidelbergensis (e) H. neanderthalensis 7. Some scientists now think that fossils identified as Homo erectus represent which two different species? (a) H. habilis and H. erectus (b) H. ergaster and H. erectus (c) H. heidelbergensis and H. ergaster

(d) H. neanderthalensis and H. erectus (e) H. neanderthalensis and H. sapiens 8. Archaic humans appeared as early as _____ years ago. (a) 5 million (b) 1.2 mya (c) 250,000 (d) 100,000 (e) 5000 9. _____ were an early group of humans with short, sturdy builds and heavy supraorbital ridges that lived throughout Europe and western Asia from about 250,000 to 30,000 years ago. (a) Australopithecines (b) Dryopithecines (c) Archaic humans (d) Neandertals (e) Cro-Magnons 10. The modern human skull lacks (a) small canines (b) a foramen magnum centered in the base of the skull (c) pronounced supraorbital ridges (d) a U-shaped arrangement of teeth on the jaw (e) a large cranium (braincase) 11. The comparison of genetic material from individuals of regional populations of humans, used to help unravel the origin and migration of modern humans, is known as (a) paleoarchaeology (b) cultural anthropology (c) molecular anthropology (d) cytogenetics (e) genetic dimorphism 12. Place the following hominins in chronological order of appearance in the fossil record, beginning with the earliest: 1. H. ergaster 2. H. sapiens 3. H. neanderthalensis 4. H. erectus (a) 1, 4, 2, 3 (b) 4, 1, 2, 3 (c) 4, 1, 3, 2 (d) 3, 1, 4, 2 (e) 1, 4, 3, 2

CRITICAL THINKING 1. If you were evaluating whether other early humans exterminated the Neandertals, what kinds of archaeological evidence might you look for? 2. The remains of H. sapiens have been found in southern Europe alongside reindeer bones, but reindeer currently exist only in northern Europe and Asia. Explain the apparent discrepancy. 3. EVOLUTION LINK. What was the common ancestor of chimpanzees and humans—a chimpanzee, a human, or neither? Explain your answer. 4. EVOLUTION LINK. Some scientists say that modern medicine and better sanitation are slowing down or altering the course of human evolution in highly developed countries today. As a result, evolution in humans today has gone from survival of the fittest to survival of almost everyone. Do you think this is a valid idea? Why or why not? 5. ANALYZING DATA. If you were to add H. floresiensis to Figure 22-9, where would you put it? Justify your answer. 6. ANALYZING DATA. Using the information in Figure 22-9, draw a simple cladogram that represents a reasonable hypothesis of human evolution at the genus level. (Hint: Your diagram will have three branches.)

7. ANALYZING DATA. The four chromosomes depicted below are Y chromosomes from men in different parts of the world. The purple bands represent specific genes that are identical in all four men. The red bands represent genetic mutations (M) that appeared at different times; M168 appeared about 50,000 years ago, M9 about 40,000 years ago, and M3 about 10,000 years ago. Based on what you have learned in this chapter, identify the geographic source of men carrying each of these chromosomes: African, Eurasian, Amerindian, and first migrants out of Africa. (You may wish to refer back to Figure 22-14.)

M168

M168 M9

M168 M9 M3

Additional questions are available in CengageNOW at www.cengage.com/ login.

23

Understanding Diversity: Systematics

AP Images

Pink iguana. Based on genetic analysis, this pink iguana was identified as a new species in 2009. The iguana inhabits the Wolf volcano on Isabela, one of the Galapagos Islands.

KEY CONCEPTS

T

23.1 Traditionally, biologists have named organisms using

described. Biologists estimate that to date they have identified less than

a binomial system (in which each species has a genus name followed by a specific epithet) and have classified organisms in taxonomic categories arranged in a hierarchy from most inclusive (domain) to least inclusive (species).

10% of bacteria, about 10% of fungi, only about 2% of nematode (round-

23.2 Based on current knowledge, the three main branches of the tree of life are the three domains: Bacteria, Archaea, and Eukarya.

he total number of living species is estimated to be between 4 million and 100 million, but only about 1.8 million species have been

worm) species, and less than 20% of insect species. Many regions on our planet have not yet been carefully inventoried, including the forest canopy, soil, and the deep regions of the ocean. Biologists discover thousands of new species each year. For example, in 2009, researchers described a pink iguana that inhabits the Galapagos

23.3 Modern systematics seeks to reconstruct phylogeny (evolutionary history of a group of species) based on common ancestry inferred from shared characters (characteristics), including structural, developmental, behavioral, and molecular similarities, as well as from fossil evidence.

Islands as a newly identified species (see photo). The research team per-

23.4 Cladograms (a type of phylogenetic tree) are diagrams of hypothetical evolutionary relationships; they are constructed by analyzing shared derived characters.

are referred to as biological diversity, or biodiversity. The totality of life

formed genetic analyses indicating that this iguana split off from other iguanas about five million years ago. The variety of living organisms and the ecosystems they are part of on Earth represents our biological heritage, and the quality of life for all

organisms depends on the health and balance of this worldwide web of life-forms. For example, we depend on organisms to maintain the life-sustaining composition of gases in the atmosphere, form soil, break down wastes, recycle nutrients, and provide food for one another. We humans exploit many species for economic benefit. One very anthropocentric (human-centered) example of the practical importance of biodiversity is that more than 40% of the prescriptions that pharmacists dispense in the United States derive from living organisms. Investigators are just learning how to effectively screen organisms for potential drugs. Unfortunately, human activity is seriously reducing biodiversity, and species are becoming extinct faster than researchers can study them. (See the discussion on declining biodiversity in Chapter 57.) Alarmed by these extinctions, biologists are mobilizing to more rapidly identify new species and to conserve biodiversity.

23.1 CLASSIFYING ORGANISMS ■ ■ LEARNING OBJECTIVES 1 2

State two justifications for the use of scientific names and classifications of organisms. Describe the binomial system of naming organisms and arrange the Linnaean categories in hierarchical fashion from most inclusive to least inclusive.

The scientific study of the diversity of organisms and their evolutionary relationships is called systematics. Recall from earlier chapters (for example, see Chapter 18) that evolution is the accumulation of inherited changes within populations over time. An important aspect of systematics is taxonomy, the science of naming, describing, and classifying organisms. In biology, the term classification means arranging organisms into groups based on similarities that reflect evolutionary relationships among lineages.

Many biologists agree that describing and classifying all the surviving species of the world should be a major scientific goal of the 21st century. Important steps toward this goal are being taken by such international research endeavors as the Catalogue of Life and the Encyclopedia of Life. The Catalogue of Life is the work of a collaborative international research team that manages a comprehensive global database made available on the Internet. The Catalogue of Life categorizes and lists all of the world’s known species. The Encyclopedia of Life (EOL), a project launched in 2007, is an electronic database that will include a home page for each species. Data will include alternative names for each organism, classification, habitat, distribution, and diet. The Convention on Biological Diversity is an international treaty made up of 190 countries committed to reducing the “rate of loss of biodiversity.” This convention is committed to developing strategies for the conservation and sustainable use of Earth’s biological diversity. In this chapter, we learn more about evolution and information transfer, two of the basic themes of life introduced in Chapter 1. We explore some of the approaches and methods used by biologists to classify organisms and to infer their evolutionary history and relationships. Information comes from many sources, including fossils, biogeography, patterns of development, and structural and behavioral characteristics. Advances in DNA sequencing have generated huge amounts of data about molecular similarities and differences. Systematists organize information about organisms they are studying into large databases and manipulate them using computer algorithms to generate hypotheses regarding evolutionary relationships. From these data, biologists are constructing a “tree of life.” As more sophisticated molecular methods and discovery of new fossils provide new data, old data are reinterpreted. As we gain greater understanding of how organisms are related, we revise the way we classify organisms, and the structure of the tree of life changes.

Organisms are named using a binomial system Because there are millions of kinds of organisms, scientists need a system for accurately identifying them. Imagine that you were going to develop a classification system. How would you use what you already know about living things to assign them to categories? Would you place insects, bats, and birds in one category because they all have wings and fly? And would you, perhaps, place squid, whales, fish, penguins, and Olympic backstroke champions in another category because they all swim? Or would you classify organisms according to a culinary scheme, placing lobsters and tuna in the same part of the menu, perhaps identifying them as “seafood”? Any of these schemes might be valid, depending on your purpose. Similar methods have been used throughout history. St. Augustine in the 4th century classified animals as useful, harmful, or superfluous—to humans. During the Renaissance, scholars began to develop categories based on the characteristics of the organisms themselves. Of the many classification systems that were developed, the one designed by Carolus Linnaeus in the mid-18th century has survived with some modification to the present day. Linnaeus grouped organisms according to their similarities, mainly structural similarities. Before the mid-18th century, each species had a lengthy descriptive name, sometimes consisting of ten or more Latin words! Linnaeus simplified scientific classification, developing a binomial system of nomenclature in which each species is assigned a unique two-part name. The first part of a binomial scientific name is a noun that designates the genus (pl., genera), and the second part, an adjective modifying that noun, is called the specific epithet. The genus name is always capitalized, whereas the specific epithet is usually not. Both names are underlined or italicized. The genus, or generic, name can be used alone to designate all species in the genus (for example, the genus Quercus includes all oak species). Note that the specific epithet alone is not the species’s name. In fact, the same specific epithet can be used as the second name of species in different genera. For example, Quercus alba is the scientific species name for the white oak, and Salix alba is the

species name for the white willow (alba comes from a Latin word meaning “white”). Thus, both parts of the name must be used to accurately identify the species. The specific epithet is never used alone; it must always follow the full or abbreviated genus name, for example, Quercus alba or Q. alba. Scientific names are generally derived from Greek or Latin roots or from Latinized versions of the names of persons, places, or characteristics. For example, the generic name for the bacterium Escherichia coli is based on the name of the scientist, Theodor Escherich, who first described it. The specific epithet coli reminds us that E. coli lives in the colon (large intestine). Most areas of biology depend on scientific names and classification. For example, to study the effects of pollution on an aquatic community, biologists must be able to accurately record the relative numbers of each type of organism present and changes in their populations over time. This requires that all investigators identify each species accurately by name. Scientific names permit biology to be a truly international science. Even though the common names of an organism may vary in different locations and languages, an organism can be universally identified by its scientific name. A researcher in Puerto Rico knows exactly which organisms were used in a study published by a Russian scientist and therefore can repeat or extend the Russian’s experiments using the same species.

TABLE 23-1 Domain

Eukarya Organisms that have nuclei and other membraneenclosed organelles

Kingdom

Plantae Terrestrial, multicellular, photosynthetic organisms

Phylum

Anthophyta Vascular plants with flowers, fruits, and seeds

Class

Monocotyledones Monocots: Flowering plants with one seed leaf (cotyledon) and flower parts in threes

Order

Commelinales Monocots with reduced flower parts, elongated leaves, and dry 1-seeded fruits

Family

Poaceae Grasses with hollow stems; fruit is a grain; and abundant endosperm in seed

Genus

Zea Tall annual grass with separate female and male flowers

Species

Zea mays Corn

Each taxonomic level is more general than the one below it Linnaeus devised a system for assigning species to a hierarchy of increasingly broader groups. As you move up the hierarchy, each group is more inclusive, that is, it includes the groups below it. When he set up his system, Linnaeus did not have a theory of evolution in mind. Nor did he have any idea of the vast number of extant (living) and extinct organisms that would later be discovered. Yet his system has proved remarkably flexible and adaptable to new biological knowledge and theory. Very few other 18th-century inventions survive today in a form that would still be recognizable to their originators. The range of taxonomic categories from species to domain forms a hierarchy (FIG. 23-1 and TABLE 23-1). Closely related species are assigned to the same genus, and closely related genera are grouped in a single family. Families are grouped into orders, orders into classes, classes into phyla, phyla into kingdoms (some biologists use the term division for this category in plants), and kingdoms into domains. A species is considered a true biological entity, but taxonomic categories above the species level, at least until recently, have been artificial constructs used to conveniently catalogue the diverse life-forms on Earth. As we will discuss in this chapter, the modern systematists work to validate taxonomic groups. A taxon (pl., taxa) is a formal grouping of organisms at any given level, such as species, genus, or phylum. For example, class Mammalia is a taxon that includes many different orders. A taxon can be separated into subgroupings, such as subphyla or superclasses. Subphylum Vertebrata, a subgroup of phylum Chordata, is a taxon that contains several classes, including Amphibia and Mammalia.

Classification of Corn

Review ■ ■

■

What are the key features of the binomial system of nomenclature? What are the advantages to using scientific, rather than common, names for organisms? Classify the domestic cat, using taxa from domain to species.

23.2 DETERMINING THE MAJOR BRANCHES IN THE TREE OF LIFE ■ ■ LEARNING OBJECTIVES 3

4

Describe the three domains and argue for and against classifying organisms in the domains. (Compare this approach with using kingdoms as the main rank of classification.) Interpret a cladogram, describing the meaning of its specific nodes and branches.

The history of taxonomy at the kingdom and domain levels is a good example of the process of science. In this section we will review some highlights of this process.

Systematics is an evolving science From the time of Aristotle to the mid-19th century, biologists divided organisms into two kingdoms: Plantae and Animalia. After the development of microscopes, it became increasingly obvious that many organisms could not be easily assigned to either the

DOMAIN Eukarya

KINGDOM Animalia

PHYLUM Chordata

CLASS Mammalia

ORDER Carnivora

FAMILY Felidae

GENUS Felis

SPECIES Felis catus

▲

FIGURE 23-1 The principal categories used in classification

The domestic cat (Felis catus) is classified here to illustrate the hierarchical organization of the Linnaean taxonomic system. Each level is more inclusive than the one below it, meaning that it includes more groups of organisms.

plant or the animal kingdom. For example, the unicellular organism Euglena has been classified at various times in the plant kingdom and in the animal kingdom, but did not really fit in either kingdom. Euglena carries on photosynthesis in the light, but in the dark, it uses its flagellum to move about in search of food (see Fig. 26-5). In 1866, a German biologist, Ernst Haeckel, proposed that a third kingdom, Protista, be established to accommodate bacteria and other microorganisms. However, biologists largely ignored this kingdom for about a hundred years. In 1937, the French marine biologist Edouard Chatton suggested the term procariotique (“before nucleus”) to describe bacteria and the term eucariotique (“true nucleus”) to describe all other cells. This dichotomy between prokaryotes and eukaryotes is now universally accepted by biologists as a fundamental evolutionary divergence. In the 1960s, advances in electron microscopy and biochemical techniques revealed further cell differences that inspired many new proposals for classifying organisms. In 1969, R. H. Whittaker proposed a five-kingdom classification based mainly on cell structure and the way that organisms derive nutrition from their environment. (Although Whittaker is generally credited, the concept of five kingdoms was actually first proposed in 1949 by T. L. Jahn and F. Jahn in their book How to Know the Protozoa.) Whittaker suggested that the fungi (which include the mushrooms, molds, and yeasts) be removed from the plant kingdom and classified in their own kingdom, Fungi. After all, fungi are not photosynthetic and must absorb nutrients produced by other organisms. Fungi also differ from plants in the composition of their cell walls, in their body structures, and in their modes of reproduction. Kingdom Prokaryotae was established to accommodate the bacteria, which are fundamentally different from all other organisms in that they do not have distinct nuclei and other membranous organelles and do not undergo mitotic division (see Fig. 4-6). In the late 1970s, Carl Woese (pronounced “woes”), of the University of Illinois, and his colleagues began to study the evolutionary relationships among organisms by analyzing their 16S rRNA. Using sequence analysis, Woese used variations in this universal molecule to challenge the long-held view that all prokaryotes were closely related and very similar to one another. He showed that there are two fundamentally different groups of prokaryotes, archaea and bacteria. Woese argued that the prokaryotes account for two of the three major branches of organisms. Woese’s hypothesis gained support in 1996 when Carol J. Bult of the Institute for Genomic Research in Rockville, Maryland, reported in the journal Science that she and her colleagues had sequenced the complete genome of a methane-producing archaea, Methanococcus jannaschii. When these researchers compared gene sequences with those of two previously sequenced bacteria, they found that fewer than half of the genes matched. Gene sequencing indicates that the archaea have a combination of bacteria-like and eukaryote-like genes. Biologists have identified other important differences between bacteria and archaea. For example, bacteria are characterized by the presence of a compound called peptidoglycan in their cell walls, whereas this compound is not present in

archaea. Based on molecular evidence, biologists now divide the prokaryotes into two major groups: Bacteria and Archaea. Kingdom Protista has had an interesting history. Biologists placed Euglena, along with other unicellular organisms traditionally referred to as protozoa, in kingdom Protista. At various times, they have also assigned algae (including multicellular forms), water molds, and slime molds to kingdom Protista. Thus, this kingdom became a diverse group of mainly unicellular, mainly aquatic eukaryotic organisms. During the past few years, systematists have established that the protist groups did not descend from one recent common ancestor. As we will discuss in Chapter 26, many biologists have abandoned kingdom Protista (as well as a kingdom classification for plants and animals) and now assign the eukaryotes to five “supergroups” based on molecular data.

The three domains form the three main branches of the tree of life Based on fundamental molecular differences among the bacteria, archaea, and eukaryotes, biologists now classify organisms in three domains: Archaea, Bacteria, and Eukarya (eukaryotes). Systematists have inferred that the three domains are the three main branches of the tree of life (FIG. 23-2). In Chapter 25 we will discuss characters that distinguish these domains. As we will discuss in Chapter 24, the viruses are a special case and are not classified in any of the three domains.

KEY POINT

Domain Bacteria

Biologists classify organisms in three major categories called domains. Domain Archaea

Domain Eukarya

Common ancestor of all living organisms

FIGURE 23-2 Animated The three domains This cladogram, a type of evolutionary tree, illustrates the evolutionary relationships among organisms in the three domains. Each branch portrays a clade, a group of organisms with a recent common ancestor. Each node (circle) represents the point when two groups diverged from each other.

Some biologists are moving away from Linnaean categories Most biologists now classify organisms into the three domains. Some continue to classify organisms into kingdoms: Bacteria, Archaea, Fungi, Plantae, and Animalia, but many systematists no longer recognize “kingdom Protista.” (See TABLE 23-2.) Instead, they assign protists to a number of “supergroups” that more accurately reflect evolutionary relationships. Many systematists argue that the traditional hierarchical Linnaean categories are limiting and do not fit well with recent findings. They maintain that using kingdoms, classes, and other ranks are cumbersome and that classification using these ranks must be frequently modified as researchers gather more data. The former kingdom Protista is a good example of the need to continuously re-evaluate and change classification. Some systematists prefer classifying organisms into clades. A clade is defined as a group of organisms that share characters (characteristics) inherited from a common ancestor. These systematists prefer to identify clades and classify them in one of the three domains, rather than using the traditional hierarchical system. Some biologists use an approach to classification known as PhyloCode, in which organisms are grouped into clades based

TABLE 23-2

on evolutionary relationships. A species is defined as a segment of a population lineage. In this system, taxonomic ranks are not required. Advocates of PhyloCode argue that this system avoids the problem of changing names when discoveries of new species require shifts in ranks. The system also makes it easier to name clades one at a time as they are discovered. In this edition of Biology, we classify organisms using the three domains and also classify them using traditional taxonomy. We remind our readers frequently that the traditional ranks are categories developed for convenience, and classification changes as new data are analyzed. Taxonomists must reclassify organisms as they adjust hypotheses about relationships among organisms.

Phylogenetic trees show hypothesized evolutionary relationships Systematists use phylogenetic trees to graphically represent hypothesized evolutionary relationships among organisms that have a common ancestor. In Figure 23-2 we use a phylogenetic tree to illustrate the relationship of the three domains, and the phylogenetic tree in FIGURE 23-3 illustrates the main branches that make up each of the three domains. This tree is one of the current versions of the tree of life.

Domains and Kingdoms

Domain

Kingdom

Characteristics

Ecological Role and Comments

Bacteria

Bacteria

Prokaryotes (lack distinct nuclei and other membranous organelles); unicellular; microscopic; cell walls generally composed of peptidoglycan.

Most are decomposers; some parasitic (and pathogenic); some chemosynthetic autotrophs; some photosynthetic; important in recycling nitrogen and other elements; some used in industrial processes.

Archaea

Archaea

Prokaryotes; unicellular; microscopic; peptidoglycan absent in cell walls; differ biochemically from bacteria.

Methanogens are anaerobes that inhabit sewage, swamps, and animal digestive tracts; extreme halophiles inhabit salty environments; extreme thermophiles inhabit hot, sometimes acidic environments.

Protists formerly classified in kingdom Protista; now assigned to a number of “supergroups”

Eukaryotes; mainly unicellular or simple multicellular.

Protozoa are an important part of zooplankton. Algae are important producers, especially in marine and freshwater ecosystems; important oxygen source. Some protists cause diseases, for example, malaria.

Plantae

Eukaryotes; multicellular; photosynthetic; possess multicellular reproductive organs; alternation of generations; cell walls of cellulose.

Terrestrial biosphere depends on plants in their role as primary producers; important source of oxygen in Earth’s atmosphere.

Fungi

Eukaryotes; heterotrophic; absorb nutrients; do not photosynthesize; body composed of threadlike hyphae that form tangled masses that infiltrate food or habitat; cell walls of chitin.

Decomposers; some parasitic (and pathogenic); some form important symbiotic relationships with plant roots (mycorrhizae) or algae (lichens); some used as food; yeast used in making bread and alcoholic beverages; some used to make industrial chemicals or antibiotics; responsible for much spoilage and crop loss.

Animalia

Eukaryotes; multicellular heterotrophs; many exhibit tissue differentiation and complex organ systems; most able to move about by muscular contraction; nervous tissue coordinates responses to stimuli.

Consumers; some specialized as herbivores, carnivores, or detritus feeders.

Eukarya*

⎛ ⎜ ⎜ ⎜ ⎜ ⎜ ⎜ ⎜ ⎜ ⎜ ⎜ ⎨ ⎜ ⎜ ⎜ ⎜ ⎜ ⎜ ⎜ ⎜ ⎜ ⎝

*For another approach to classification of eukarya, see Figure 23-3.

Domain Archaea

Domain Eukarya

Animals

Unikonts

Fungi

Land plants

Green algae

Red algae

Rhizarians

Chromalveolates

Excavates

Crenarchaeota

Euryarchaeota

Actinobacteria

Spirochetes

Chlamydias

Gram-positives

Cyanobacteria

Proteobacteria

Archaeplastids

Choanoflagellates

Domain Bacteria

Most recent common ancestor of all living things

FIGURE 23-3 The tree of life, a work in progress Some major branches of each of the three domains are shown. As new data are analyzed, the relationships of some of the branches will change.

The type of phylogenetic tree we use in this book is called a cladogram. Each branch in a cladogram represents a clade, a group of organisms with a common ancestor. Each branching point, referred to as a node (depicted by a circle), represents the divergence, or splitting, of two or more new groups from a common ancestor. Thus, the node represents the most recent common ancestor of each clade depicted by the branches. In this way, a cladogram uses the positions of branch points to illustrate the hypothesized evolutionary relationships among taxa. Each branch represents one or more characteristics shared by the clade, but not found in the ancestors of the clade. (Shared derived characters are discussed in the next section). These shared characteristics can be indicated by labels or by bars across the branches. Cladograms are rooted if the most recent common ancestor is known. The root, or node at the base of the cladogram, represents the most recent common ancestor of all of the clades depicted in the tree. We will discuss more about constructing cladograms as we proceed through this chapter. Although a cladogram depicts the evolutionary relationships of each group through time, the lengths of the branches do not tell us when a particular species evolved. Other types of phylogenetic trees, for example, phylograms, can be constructed to indicate time, or rate of evolution, as well as to indicate the relationships among

taxa. In a phylogram, the length of the branches is proportional to the amount of inferred change in characteristics.

Systematists continue to consider other hypotheses When we think of how organisms acquire their genes, we think of vertical gene transfer in which genes are transmitted from parent to offspring within the same species. As biologists continue to investigate the origin and relationships of the domains, they have discovered that evolution is not always linear. During the course of evolution, genes not only were passed down “vertically” from one generation to the next but were also exchanged laterally. Such gene swapping between organisms in one taxon and unrelated organisms in another taxon is called horizontal gene transfer, or lateral gene transfer. In this process, genes move from one species to another species in the same generation. Horizontal gene transfer can occur in several ways, such as by exchange of DNA among different populations or species of bacteria, or by interbreeding between closely related groups. In eukaryotes, horizontal gene transfer has occurred by endosymbiosis, a process in which one organism lives inside the cell of another organism, and the two become a single functional organism. In fact,

eukaryotic cells most likely evolved from prokaryotic cells that lived symbiotically one within another. Recall that mitochondria and chloroplasts are thought to have originated from prokaryotes that lived as endosymbionts inside other cells (see Chapter 21). Based on what is now known about horizontal gene transfer, some systematists argue that the common ancestor of all living things may have been a community of species that traded their genes. Some systematists suggest that there is no simple tree of life and have proposed alternative hypotheses. One group hypothesizes a complex bush with many connecting branches (FIG. 23-4a). Researchers at the University of California at Los Angeles, have re-

Crenarchaeota

Euryarchaeota

Archaea

Animalia Fungi Plantae Protists Protists Protists

Eukarya Cyanobacteria

Proteobacteria

Bacteria

cently proposed another approach—the tree of life may actually be a ring of life (FIG. 23-4b). These systematists hypothesize, based on their analysis of hundreds of genes, that horizontal gene transfer between bacteria and archaea gave rise to the eukaryotes. Horizontal gene transfer appears to be a continuous process among members of the three domains. The evolution of systematics reflects the creative and dynamic process of science. Systematists are very responsive to new data, and consequently, classification of organisms at all levels is a continuously changing process. Although many biologists are moving away from a hierarchical system, some suggest adding new groups so that the classical approach keeps up with new findings. Whatever approach is used, the goal of systematics is to base classification on evolutionary change and relationships.

Review ■

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(a) The three-domain approach drawn to show horizontal gene transfer as a continuous process between domains and also among groups within each domain. Only some of the branches are labeled. The diagonal brown arrow represents mitochondrial endosymbiosis; the diagonal green arrow represents chloroplast endosymbiosis.

23.3 RECONSTRUCTING EVOLUTIONARY HISTORY ■ ■ LEARNING OBJECTIVES 5 6 7

Protists Plants Fungi Animals

First eukaryote Bacteria

Archaea

(b) The ring of life is based on the hypothesis that lateral gene transfer between bacteria and archaea gave rise to the eukaryotes.

FIGURE 23-4 Other ways to represent the tree of life (Part a based on Doolittle, W. F., Science, Vol. 284, pp. 2124–2128, 1999; part b based on Rivera, M. C., and J. A. Lake. Nature, Vol. 431, pp. 152–155, Sept. 9, 2004.)

What are some of the defining characteristics of each of the three domains? of kingdom Fungi? In which domain and kingdom would you classify each of the following: white willow, the bacterium Escherichia coli, tapeworm, and black bread mold? What does each branch represent in a cladogram? each node?

8

Critically review the difficulties encountered in choosing taxonomic criteria. Apply the concept of shared derived characters to the classification of organisms. Describe how analyses of molecular homologies contribute to the science of systematics. Contrast monophyletic, paraphyletic, and polyphyletic taxa.

Modern taxonomy is based on evolution. The goal is to reconstruct phylogeny (literally, “production of phyla”), the evolutionary history of a group of organisms from a common ancestor. As systematists determine evolutionary relationships among species and between species and higher taxa, they build classifications based on common ancestry. Consequently, systematics reflects the integration of all evolutionary processes and evolutionary evidence. Systematics is at the center of how we understand and explain the nonrandom occurrence and relationships of life-forms on Earth. Once phylogenies are established, they help answer other questions in biology. For example, phylogenies help us understand evolutionary patterns that might provide clues to the origin and spread of HIV and other pathogens. Phylogenies also help biologists identify new species and predict their characteristics. Systematists may classify a new species based on specific characters that it shares with other organisms in a particular taxon. They may then infer that the new species shares other characters with organisms in that group. As you read the following sections, remember that phylogenies are testable hypotheses. They are supported or falsified by the

available data. Systematics proceeds by constantly re-evaluating data, hypotheses, and theoretical constructs. As new data are discovered and old data are reinterpreted, systematists modify their hypotheses. As a result, our understanding of how organisms are related and the way we classify organisms are continuously being revised. Systematics is a dynamic science that changes as biologists discover new species and use ever more sophisticated techniques to investigate the evolutionary relationships among organisms. Recall that a population is made up of all the individuals of the same species that live in a particular area. A population has a dimension in space—its geographic range—and also a dimension in time. Each population extends backward in time. Somewhat like branches of a tree, a population may diverge from other populations enough to become a new species (which may be depicted by a new tree branch; see Chapters 19 and 20). Species have various degrees of evolutionary relationships with one another, depending on the degree of genetic divergence since their populations branched from a common ancestor.

Homologous structures are important in determining evolutionary relationships Just how to determine phylogeny and how to group species into higher taxa—genera, families, orders, classes, or phyla—can be difficult decisions. Biologists base their judgments about the degree of relationship on the extent of similarity among species. They examine structural, physiological, developmental, behavioral, and molecular traits, as well as fossil evidence. When examining these traits, they look for homologies among different organisms. Recall from Chapter 18 that homology refers to the presence, in two or more species, of a structure derived from a recent common ancestor. For example, the bones in the wing of a bat, the paw of a dog, and the hand of a human are homologous. As we will discuss, identifying homologous traits is very important in inferring phylogeny. It is often difficult to determine whether traits are similar as a result of homology. Although the wings of a butterfly are adapted for flight, their structure is different from bird wings, and these animals do not share a common winged ancestor. Similar structures sometimes evolve when unrelated or distantly related species become adapted to similar environmental conditions. Thus, wings may occur in two or more species not derived from a recent common ancestor. Recall from Chapter 18 that independent evolution of similar structures in distantly related organisms is known as convergent evolution. Sharks and dolphins have similar, but independently derived, body forms because they have become adapted to similar environments (aquatic) and lifestyles (predatory). Sometimes it is obvious that convergent evolution has occurred. In other cases, researchers must test their hypotheses using all available data. Another challenge in deciding on homology is reversal, in which a trait reverts to its ancestral state. A reversal removes a similarity that had evolved. A characteristic that superficially appears homologous but is actually independently acquired by convergent evolution or reversal is described as exhibiting homoplasy. Distinguishing between homology and homoplasy can be challenging.

Shared derived characters provide clues about phylogeny Determining which traits indicate evolutionary relationships is extremely important. How does a systematist interpret the significance of similarities between organisms? In making decisions about taxonomic relationships, the systematist first examines the characteristics in the largest group (such as phylum or class) of organisms being studied and interprets them as indicating the most remote common ancestry. These shared ancestral characters, or plesiomorphies, are features that were present in an ancestral species and remain present in all groups descended from that ancestor. For example, the vertebral column, present in all vertebrates, is an ancestral character for study of classes within the subphylum Vertebrata. Studying the presence or absence of the vertebral column does not help us discriminate among various classes of vertebrates (for example, between amphibians and mammals) because all individuals in these classes have a vertebral column. When two populations become separated and begin to evolve independently, some of their homologous traits change as a result of mutation, natural selection, and genetic drift. The novel traits that evolve are referred to as shared derived characters, or synapomorphies. Note that shared derived characters originate in a recent common ancestor and are present in its descendants. Species that share derived characters form a clade. Systematists use shared derived characters to identify points where groups diverged from one another. A trait viewed as a derived character in a more inclusive (broader) taxon may also be considered an ancestral character in a less inclusive (narrower) taxon. More recent common ancestry is indicated by classification into less and less inclusive taxonomic groups with more and more specific shared derived characters. For example, the three small bones in the middle ear are useful in identifying a branch point between reptiles and mammals. The evolution of this derived character was a unique event, and only mammals have these bones. However, if we compare mammals with one another, the three ear bones are a shared ancestral character because all mammals have them. Consequently, they have no value in distinguishing among mammalian taxa. Other characters must be used to establish branch points among the mammals. If we compare dogs, goats, and dolphins (all of which are mammals), we find that dogs and goats have abundant hair whereas dolphins do not. Hair is an ancestral trait in mammals and therefore cannot be used as evidence that dogs and goats share a more recent common ancestor. In contrast, the virtual absence of hair in mature dolphins is a derived character within mammals. When we compare dogs, dolphins, and whales, we find that dolphins and whales share this derived character, providing evidence that these animals evolved from a common ancestor not shared by dogs.

Biologists carefully choose taxonomic criteria Both fishes and dolphins have streamlined body forms, but this characteristic is homoplastic and does not indicate close evolutionary relationships. In contrast, dolphins share important homologous derived characters (synapomorphies) with mammals:

later, though, birds evolved from a group of dinosaurs whose fossils bear impressions of feathers! The presence of feathers in some dinosaurs and in birds is a shared derived character that shows that they are closely related. Many biologists now classify birds as part of the reptile clade. Organisms are typically classified on the basis of a combination of traits rather than on any single trait. The significance of these combinations is determined inductively, that is, by integrating and interpreting the data. Such induction is a necessary part of the process of science. Systematists may hypothesize that all birds should have beaks, feathers, no teeth, and so forth. Then they reexamine the living world and observe whether any organisms might reasonably be called birds that do not fit the current definition of “birdness.” If not, the definition stands. If too many exceptions emerge, the definition may be modified or abandoned. Sometimes the systematist determines that an apparent exception—the bat, for instance—resembles a bird only superficially and should not be considered one. The bat has all the basic characteristics of a mammal, such as hair and mammary glands that produce milk for the young.

mammary glands, which produce milk for the young; three small bones in the middle ear; and a muscular diaphragm that helps move air into and out of the lungs. Thus, we classify the dolphin as a mammal. Although dolphins have more shared derived characters in common with humans than with fishes, some characters are shared by all three. Among these shared ancestral characters are a dorsal tubular nerve cord, and during embryonic development, a notochord (skeletal rod) and rudimentary gill slits. These shared ancestral characters (plesiomorphies) indicate a common ancestry and serve as a basis for classification. The ancestry is more remote between the dolphin and the fish than between the dolphin and the human. Therefore, although fishes, humans, and dolphins are grouped in a more inclusive taxon, the phylum Chordata, humans and dolphins are also classified in class Mammalia, a less inclusive taxon within phylum Chordata, indicating that they are more closely related. Determining which traits best illustrate evolutionary relationships can be challenging. What, for example, are the most important taxonomic characteristics of a bird? We might list feathers, beak, wings, absence of teeth, egg laying, and endothermy (an endothermic animal uses metabolic heat to maintain a constant body temperature despite variations in surrounding temperature). The duck-billed platypus and a few other mammals (members of a group called monotremes) have many of these same characteristics: beaks, absence of teeth, egg laying, and endothermy. Yet we do not classify them as birds (FIG. 23-5). No mammal, however, has feathers. Is this trait absolutely diagnostic of birds? According to the conventional taxonomic wisdom, the presence or absence of feathers determines what is and is not a bird. As we will discuss

© Jean Philippe Varin/Jacana/Photo Researchers, Inc.

Molecular homologies help clarify phylogeny

FIGURE 23-5 Is this animal a bird? A few mammals, such as the duck-billed platypus, lay eggs, have beaks, and lack teeth. However, the platypus does not have feathers, and it nourishes its young with milk secreted from mammary glands.

When a new species evolves, it does not always exhibit obvious phenotypic differences when compared to closely related species. For example, two distinct species of fruit flies may appear identical. Some of their DNA, proteins, and other molecules, however, are different. Such variations in the structure of specific macromolecules among species, just like differences in anatomical structure, result from mutations. Advances in molecular biology have provided the tools for biologists to compare the macromolecules of various organisms. In fact, organisms can now be identified by their molecular structure. In 2003, scientists at the University of Guelph in Canada proposed in the British Proceedings of the Royal Society that we identify all living things by their unique sequences of DNA or RNA, rather than by their physical structure. The molecular sequence selected can be used as a genetic marker, or barcode, to identify organisms, much like the barcodes on grocery store items. This method can be used to distinguish among species that look alike. For example, until recently the African elephant and the Indian elephant were the only two species of living elephants recognized. Using molecular methods, biologists demonstrated that there are at least two distinct species of African elephants. Each species has its own distinctive DNA barcode. Biologists can use barcodes to identify the species of an organism in any stage of life, even as an egg or seed. The science of molecular systematics focuses on molecular structure to clarify evolutionary relationships. DNA, RNA, and amino acid sequencing are used to compare the macromolecules of organisms being studied. Macromolecules that are functionally similar in two different types of organisms are considered homologous if their subunit sequence is similar. Such comparisons provide systematists with valuable information about the degree of relatedness among organisms. The more subunit sequences of two species correspond, the more closely related the species are considered to be. The number of differences in certain DNA or RNA nucleotide

Taxa are grouped based on their evolutionary relationships Most known organisms were classified by biologists before the days of phylogenetics. Modern systematists use phylogenetic analysis to test these hypothesized relationships among groups of organisms. Based increasingly on molecular data, they either

KEY EXPERIMENT QUESTION: How is the dog related to other canids? HYPOTHESIS: Analysis of molecular data from a variety of canids will clarify the evolutionary relationships of canids.

African wild dog (Lycaon pictus)

Dhole (Cuon alpinus)

Ethiopian wolf (Canis simensis)

Coyote (Canis latrans)

Gray wolf (Canis lupus)

Golden jackal (Canis aureus) Dog (Canis lupus familiaris)

Black-backed jackal (Canis mesomelas)

EXPERIMENT: Researcher Kerstin Lindblad-Toh and her colleagues sequenced the genome of the domestic dog. They compared coding regions of more than 13,000 dog genes with corresponding human and mouse genes. Then they selected 12 exons and 4 introns and sequenced them in 30 out of 34 living canids.

Side-striped jackal (Canis adustus)

sequences or in amino acid sequences in two groups of organisms may reflect how much time has passed since the groups branched from a common ancestor. (This can be true only if the molecular changes occur at a steady rate.) Thus, specific macromolecules can be used as molecular clocks. Many systematists look to ribosomal RNA structure to help determine phylogenies. All known organisms have ribosomes that function in protein synthesis, and certain ribosomal RNA nucleotide sequences have been highly conserved in evolution. Recall that the division of organisms into three domains was based, in large part, on the comparison of ribosomal RNA by Carl Woese and his research team. The ribosomes of archaea and bacteria contain three types of RNA, named in order of increasing size: 5S, 16S, and 23S. (These numbers are sedimentation coefficients, measures of relative size, used to characterize behavior of a particle when centrifuged.) The 5S and 16S RNAs have been extensively used to determine evolutionary relationships among bacteria. These rRNA gene sequences are useful because the number of base pairs is manageable and because these molecules are transcribed from highly conserved regions of DNA. Also, these gene sequences contain variable regions that are species-specific for various types of bacteria and archaea. Researchers have also used comparison of ribosomal RNA sequences to challenge the once widely accepted idea that fungi are closely related to plants. According to ribosomal RNA analysis, fungi are more closely related to animals than to plants; that is, animals and fungi share a more recent common ancestor, perhaps a flagellate. Because genes can evolve at different rates, researchers now typically compare subunit sequences from many different genes of the organisms under study. Molecular sequence data have been sampled from more than 10% of all known species. The very precise tools provided by molecular biology have put systematics on the cutting edge of biological research. Scientists at Broad’s Genome Sequencing and Analysis program at MIT and Harvard recently provided another example of applied molecular systematics (FIG. 23-6). They sequenced the genome of the domestic dog and compared coding regions of more than 13,000 dog genes with human and mouse genes. Then, they selected exons and introns and compared their nucleotide sequences in 30 living canids (members of the family Canidae to which the dog belongs). The data they gathered enabled these investigators to construct a cladogram of the phylogenetic relationships of canids. They confirmed earlier research showing that the dog’s closest relative is the gray wolf. These nucleotide data are being used to identify genes responsible for specific diseases, and can be compared to the human genome in an effort to better understand the many diseases that canids have in common with humans.

RESULTS AND CONCLUSION: Analysis of the comparisons of exon and intron nucleotide sequences allowed these researchers to build a cladogram of Canidae species. The data indicated that the dog is most closely related to the gray wolf. (Sequence divergence of nuclear exon and intron sequences is 0.04% and 0.21, respectively.) The two African jackals (branch at left) are the sister group of this clade, suggesting an African origin for the canids. Lindblad-Toh, K., et al. "Genome Sequence, Comparative Analysis and Haplotype Structure of the Domestic Dog." Nature, Vol. 438, Dec. 8, 2005.

FIGURE 23-6 Molecular phylogeny Molecular similarities indicate that the dog is closely related to the gray wolf. Note that some biologists list the scientific name for the domestic dog as Canis familiaris and others, including the Smithsonian Institution and the American Society of Mammalogists, consider the dog a subspecies of the gray wolf and list its name as Canis lupus familiaris.

confirm or modify the work of earlier biologists. Using their data, systematists construct cladograms to reflect relationships among clades. Cladograms show three types of taxonomic relationships: monophyletic, paraphyletic, and polyphyletic. A monophyletic group includes an ancestral species and all its descendants (FIG. 23-7a). It is defined by shared derived characters. Mammals, for example, have mammary glands that produce milk for the young, three small bones in the middle ear, and a muscular diaphragm. All mammals are thought to have evolved from a common ancestral mammal that had these shared derived

KEY POINT

Groups of organisms can be described as monophyletic, paraphyletic, or polyphyletic. Only monophyletic groups are considered clades. Group II

Group III

Group I 1

2

1 3

C

4

5

Common ancestor of groups 2 and 3

2

3

6

4

E Common D ancestor of groups 4, 5, and 6

E D

Common ancestor of groups 4, 5, and 6

B A

Common ancestor of all groups shown

(b) Group III is paraphyletic. It includes some, but not all of the descendants of the recent common ancestor indicated at node D.

(a) Group I and group II are monophyletic groups, or clades. Each includes a common ancestor and all its descendants.

Group IV

1

2

3

C

4

5

Most recent common ancestor of groups 2 and 3

FIGURE 23-7 Animated Evolutionary relationships In all three figures, the branch point at node A represents the common ancestor of all groups shown. Node B represents the common ancestor of groups 2 through 6. Node C represents the common ancestor of groups 2 and 3. Node D represents the common ancestor of groups 4, 5, and 6. Node E represents the common ancestor of groups 5 and 6. Common ancestry is the basis for making decisions about classification.

characters, and all descendants of this ancestor are mammals. Monophyletic taxa are called clades. They are natural groupings because they represent true evolutionary relationships and include all close relatives. Sister taxa, or sister groups, share a more recent common ancestor with one another than either taxon does with any other group shown on a cladogram. In Figure 23-7, taxa 2 and 3 are sister taxa; taxa 5 and 6 are also sister taxa. A paraphyletic group is a group that contains a common ancestor and some, but not all, of its descendants (FIG. 23-7b). Members of the group share ancestral characters. As discussed later in this chapter, the class Reptilia is paraphyletic because it does not include all descendants of the most recent common ancestor of reptiles. Birds share a recent common ancestor with reptiles. A polyphyletic group consists of several evolutionary lines that do not share the same recent common ancestor (FIG. 23-7c).

6

C

B A

5

6

E Most recent D common ancestor of groups 4, 5, and 6

B A

(c) Group IV is polyphyletic. Members of this group do not share the same recent common ancestor.

Biologists might have mistakenly classified the members of such a group together because these organisms share similar (homoplastic) features arising from convergent evolution. Systematists avoid constructing polyphyletic taxa because they are unnatural and misrepresent evolutionary relationships. Polyphyletic taxa are sometimes accepted temporarily until research provides additional data.

Review ■

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How are shared ancestral characters and shared derived characters different? Why don’t shared ancestral characters provide evidence for relationships between organisms within a taxon that has those traits? Give an example. How is molecular biology contributing to the science of systematics? Why do systematists prefer monophyletic taxa to polyphyletic taxa? Why do representations of the tree of life constantly change?

23.4 CONSTRUCTING PHYLOGENETIC TREES ■ ■ LEARNING OBJECTIVES 9 Contrast the classification of reptiles and birds by a cladist and by an evolutionary systematist.

10 Describe the construction of a cladogram by using outgroup analysis. 11 Apply the principle of parsimony to cladogram construction.

In determining the relationships among organisms, systematists use a variety of data and methods. How data are analyzed and interpreted depends on the systematist’s approach. In the phenetic approach, also known as numerical taxonomy, classification is based on the number of shared characters. This approach does not distinguish between shared ancestral characters and shared derived characters. Phenetic techniques are currently used with molecular data. For example, each amino acid in a protein can be considered a character. Then, amino acid sequences of various animals can be determined in the laboratory and compared by computer. The information about differences in amino acid sequences can be used to construct phylogenetic diagrams. Species are placed at relative distances from one another, reflecting the extent of difference in amino acid sequence. Thus, systematists using a phenetic approach define phylogenetic relationships by statistics, such as genetic distances, which summarize overall similarities among taxa based on collected data. A researcher can compare the base sequences of a given gene or coding region to arrive at an average percentage of difference between the taxa being analyzed. A computer program then compares these values and generates trees that cluster similar taxa and place divergent taxa on more distant branches. Cladistics (also known as phylogenetic systematics) is an approach in which organisms are classified based on recent common ancestry. Shared derived characters are analyzed and used to infer evolutionary relationships. Cladists emphasize common ancestry rather than phenotypic similarity as the basis for classification. They base their assessment on shared derived characters that can be structural, behavioral, physiological, or molecular. The characters must be homologous. Cladists typically do not use a hierarchy of taxa, for example, order, class, family. According to cladistics, dolphins are classified with mammals rather than with fishes because dolphins and mammals share derived characters not present in fishes. Their shared derived characters indicate that dolphins and mammals share a more recent common ancestor. Cladists use shared derived characters to reconstruct evolutionary relationships of organisms, and they express them in cladograms. Consider the grouping of mammals, lizards, snakes, crocodiles, dinosaurs, and birds. Birds, along with dinosaurs, are thought to share a recent common ancestor with modern crocodiles and alligators (node D in FIG. 23-8a). Crocodiles, dinosaurs, and birds, then, constitute a monophyletic group, or clade. Similarly, snakes and lizards form a clade that is the closest group to birds, dinosaurs, and crocodiles. Mammals make up a different clade.

Biologists who use the traditional evolutionary systematic approach use phenotypic similarity and a combination of shared ancestral characters and shared derived characters to establish evolutionary relationships and build classifications. Although most of the taxa that they currently recognize are monophyletic, evolutionary systematists recognize paraphyletic taxa as well. (Recall that paraphyletic groups include some, but not all, subgroups of organisms that share the same most recent common ancestor.) In contrast to cladists, evolutionary systematists recognize class Reptilia as a valid group containing snakes, lizards, crocodiles, dinosaurs, and turtles, even though this class is paraphyletic; it does not include all subgroups, such as birds, that evolved from the ancestral reptile (FIG. 23-8b). Evolutionary systematists assign birds to a separate class because birds have diverged markedly from the reptiles.

Outgroup analysis is used in constructing and interpreting cladograms A crucial step in most cladistic analyses is outgroup analysis, a research method for estimating which attributes are shared derived characters in a given group of organisms (FIG. 23-9). An outgroup is a taxon that is considered to have branched off earlier than the taxa under investigation, the ingroups. An ideal outgroup is the closest relative of the group being studied, its sister taxon, and has not been highly modified since its origin. Recall that sister taxa evolved from the same recent common ancestor. Systematists argue that an outgroup is likely to retain the ancestral state for characters being used in the analysis, allowing the researchers to identify the evolutionary changes leading to derived characters. To help you understand outgroup analysis, we describe the specific example illustrated in Figure 23-9. The first step in constructing a cladogram is to select the taxa, which may consist of individuals, species, genera, or other taxonomic levels. Here we use a representative group of eight chordates: the lancelet, lamprey, sunfish, frog, lizard, bear, chimpanzee, and human. The next step is to select the homologous characters to be analyzed. In our example, we use seven characters. For each character, we must define all the different conditions, or states, as they exist in our taxa. For simplicity, we consider our characters to have only two different states: present or absent. Keep in mind that many characters used in cladistics have more than two states. For example, black, brown, yellow, and red may be only a few of the many possible states for the character of hair color. The last, and often the most difficult, step in preparing the data is to organize the character states into their correct evolutionary order. For this step, we use outgroup analysis. In our example, the lancelet, a small marine chordate with a fishlike appearance, is the chosen outgroup. It belongs to a taxon that is considered to have diverged earlier than any of the other taxa under investigation but to be closely related to vertebrates; thus, the lancelet represents an approximation of the ancestral condition for vertebrates. Therefore, the character state “absent” for a particular character state such as vertebrae is the ancestral (plesiomorphic) condition, and the character state “present” is the derived condition for the characters in Figure 23-9.

KEY POINT

Many biologists prefer the cladistic approach, shown in (a), because it recognizes reptiles and birds as a monophyletic group.

C

C

G

Mammals

G

F

F

E

E

D

D

B

B

A

A

Most recent common ancestor of all groups shown

(a) Cladists classify birds and reptiles together because they have a recent common ancestor and are a monophyletic group. The cladogram shows the branching points in the evolution of the major groups of reptiles. Lizards, snakes, and crocodiles are phenotypically most similar, but crocodiles, dinosaurs, and birds are most closely related because they evolved most recently from a common ancestor (node D). Node F represents the branching of two clades of dinosaurs from a common ancestor, and the subsequent branching of birds from the dinosaurs.

Birds

Saurischian dinosaurs (extinct)

Ornithischian dinosaurs (extinct)

Flying reptiles (extinct)

Crocodiles, alligators

Lizards, snakes

Tuataras

Turtles

Mammals

Birds

Saurischian dinosaurs (extinct)

Ornithischian dinosaurs (extinct)

REPTILES

Flying reptiles (extinct)

Crocodiles, alligators

Lizards, snakes

Tuataras

Turtles

REPTILES

Most recent common ancestor of all groups shown

(b) Evolutionary systematists recognize the common ancestry of birds and reptiles, but assign them to different classes because of their many unique adaptations. In this approach, reptiles are a paraphyletic group.

FIGURE 23-8 Two approaches to the classification of reptiles and birds

A cladogram is constructed by considering shared derived characters Our objective is to construct a cladogram that requires the fewest number of evolutionary changes in the characters. Taxa are grouped by the presence of shared derived characters. To form a valid monophyletic group, all members must share at least one derived character. Membership in a clade cannot be established by shared ancestral characters (plesiomorphies). In our example, notice that all taxa, except the outgroup lancelet, have vertebrae. We can therefore conclude that these seven vertebrate taxa form a valid clade. Next, among the seven vertebrate taxa, note that jaws are present in all groups except lampreys (a group of jawless vertebrates). Using these data, we construct a preliminary cladogram (see FIG. 23-9a). Recall that the root, or base, of the cladogram represents the common ancestor for all taxa being analyzed. In Figure 23-9a, node A represents the common chordate ancestor from which the outgroup (lancelet) and the seven vertebrate taxa evolved. Similarly, node B represents the common ancestor of the vertebrates; and node C, the common ancestor of the jawed vertebrates. Con-

tinuing with this procedure, notice that among the six-jawed taxa, all but sunfish are tetrapods (animals with four limbs) (see FIG. 23-9b). Among the five tetrapods, all but frogs have amniotic eggs in which the embryo is surrounded by a fluid-filled sac called an “amnion” (see FIG. 23-9c). The branching process is continued, using the data in the table in the top center of the figure, until all clades are established (see FIG. 23-9d).

In a cladogram each branch point represents a major evolutionary step Notice that humans and chimpanzees share a recent common ancestor at node G in Figure 23-9d. This hypothesis is supported by the derived characters they share (synapomorphies). In the same way, bears are more closely related to the human–chimpanzee clade than to any other clade considered in our example, as indicated by the common ancestor at node F. Note that shared derived characters are nested. As you trace the tree from its root to its tips, each branch reflects the addition of one or more shared derived characters. When we compare the nodes, the order of divergence (branching) is indicated by relative

RESEARCH METHOD

Constructing a cladogram using outgroup analysis

Outgroup analysis is used to organize character states into their evolutionary order, that is, to reconstruct phylogeny. = Absent

P

= Present

CHARACTERS

Lancelet (outgroup)

A

A

A

A

A

A

A

Lamprey

P

A

A

A

A

A

A

Sunfish

P

P

A

A

A

A

A

Frog

P

P

P

A

A

A

A

Lizard

P

P

P

P

A

A

A

Bear

P

P

P

P

P

A

A

Chimpanzee

P

P

P

P

P

P

A

Human

P

P

P

P

P

P

P

TAXA

Tetrapod limbs

Node B

Node C

Node A

Node C

Common jawed vertebrate ancestor

Node B Node A

Common vertebrate ancestor

Common chordate ancestor

Human

Bear

Frog

Lizard

Present

Sunfish

Lamprey

Lancelet

Human

Chimpanzee

Absent

Bear

Lizard

Frog

Sunfish

Present

Lamprey

Lancelet

Absent

Chimpanzee

Jaws

Tetrapod (4 limbs) Amniotic egg Mammary glands Opposable thumb Upright posture

Jaws

A

Vertebrae (backbones)

Why Is It Used?

Common tetrapod ancestor Common jawed vertebrate ancestor

Common vertebrate ancestor

Common chordate ancestor

(a) All of the seven vertebrate taxa shown here have jaws, except the lamprey. Jaws are a shared derived character for these six taxa.

(b) Tetrapod limbs are a shared derived character for all vertebrate taxa shown here except the lamprey and sunfish. 2

Amniotic egg

Opposable thumb

Node G

Node F

Common amniote ancestor Common tetrapod ancestor Common jawed vertebrate ancestor Common vertebrate ancestor Node D

Node C Node B Node A

Common chordate ancestor

(c) The taxa represented by lizard, bear, chimpanzee, and human are amniotes. They share the derived character amniotic egg.

How Is It Done?

Node C Node B

Human

Chimpanzee

Bear

Lizard

Present

Frog

Sunfish

Lamprey

Lancelet

Human

Absent Chimpanzee

Bear

Frog

Lizard

Present

Sunfish

Lamprey

Lancelet

Absent

Common primate ancestor

Node Common mammal E ancestor Node D Common amniote ancestor

Common tetrapod ancestor Common jawed vertebrate ancestor

Node A

Common vertebrate ancestor Common chordate ancestor

(d) Of the vertebrate taxa shown here, only the chimpanzee and human share the derived character opposable thumb.

1 Select the taxa of interest and the homologous characters to be analyzed. 2 Define the possible conditions, for example, present (P) or absent (A). 3 Here, the lancelet is selected as the outgroup, a taxon that diverged earlier than any of the other taxa being considered. The lancelet represents an approximation of the ancestral condition. 4 Refer to the table as you follow steps (a) through (d) to build a cladogram.

FIGURE 23-9 Animated Constructing a cladogram

Opposable thumb

Opposable thumb

Node E

Node D Node C Node B Node A

Common primate ancestor

Node E

Common mammal ancestor

Node D

Common amniote ancestor

Common tetrapod ancestor

Node C Node B

Common jawed vertebrate ancestor Common vertebrate ancestor

Node A

Common chordate ancestor

(a) This cladogram, the style used in this book, has diagonal branches.

When systematists consider the evolutionary relationships of organisms, they must choose between multiple, competing clado-

(b) This is another way to depict the same relationships shown in (a). Opposable thumb Lancelet (outgroup)

Common chordate ancestor

Lamprey Sunfish

Vertebrae Frog

Jaws

Lizard

Tetrapod

Bear

Amniotic egg Mammary glands Opposable thumb

Chimpanzee Human

Upright posture

(c) This cladogram has rectangular branches and is rotated 90 degrees. However, it illustrates the same relationships as the cladograms in (a) and (b).

FIGURE 23-10 Cladogram styles These three cladograms look very different, but they depict the same relationships. Remember that the relationships are depicted by which groups branch at the nodes. The length of the branches is not important.

grams. How do they choose the most accurate branching pattern? The most common criterion is the principle of parsimony—they choose the simplest explanation to interpret the data. Parsimony, a guiding principle in many areas of research, is based on the experience that the simplest explanation is probably the correct one. Applied to choice of cladograms, parsimony requires that the clad-

Present

Systematists use the principles of parsimony and maximum likelihood to make decisions

Common chordate ancestor

Absent

distance from the base of the diagram. The farther a node is located from the base of the cladogram, the more recently the group diverged. In our example, node G represents the most recent divergence, and node A represents the most ancient divergence. Thus, in our example, humans are closely related to chimpanzees (through node G) but more distantly related to bears (through node F). Therefore, classical biologists assign humans and chimpanzees to a less inclusive taxon (order Primates), whereas humans, chimpanzees, and bears are assigned to a broader, more inclusive taxon (class Mammalia). In addition, the cladogram reveals that lizards are more closely related to the mammal clade than to frog, sunfish, or any other clade. Can you explain why? Two important concepts guide interpretation of cladograms. First, the relationships among taxa are determined only by tracing along the branches back to the most recent common ancestor and not by the relative placement of the branches along the horizontal axis. It is possible to represent the same relationships with many different types of branching diagrams (FIG. 23-10). Note that all three cladograms in this figure are equivalent. (Verify this by comparing the nodes and by checking the relationships described earlier.) Thus, a tree can be rotated around its nodes. The branching relationships are important, not the order of the taxa or the size of the branches. A second important concept is that the cladogram tells us which taxa shared a common ancestor and how recently they shared a common ancestor. The ancestor itself remains unspecified. The cladogram does not establish direct ancestor–descendant relationships among taxa. In other words, a cladogram does not suggest that a taxon gave rise to any other taxon.

Lamprey

Node F

Frog

Node G

Absent

Bear

Human

Lizard

Sunfish

Lancelet

Present Chimpanzee

Node G

Node F

Absent

Human

Chimpanzee

Bear

Lizard

Present

Frog

Sunfish

Lamprey

Lancelet

Absent

RESEARCH METHOD

A B C

(a) Hypothesis 1: Dolphins and bony fishes are close relatives.

Dolphins

Terrestrial mammals

Reptiles

Amphibians

A B C

A B C

Common ancestor

How Is It Done?

Bony fishes

Sharks

Jawless fishes

Terrestrial mammals

Reptiles

Amphibians

Dolphins

Cladogram construction is used to choose between alternative hypotheses of evolutionary relationships by selecting the most accurate branching pattern.

Bony fishes

Sharks

Jawless fishes

Why Is It Used?

Applying the principle of parsimony to cladogram construction

Common ancestor

(b) Hypothesis 2: Dolphins and terrestrial mammals are close relatives.

Systematists choose the simplest explanation to interpret the data. 1 Draw the possible cladograms. 2 Examine the relevant characters. In this example, character A = hair; B = mammary glands; C = middle ear bones. 3 Note that in the first cladogram (hypothesis 1), all three characters examined had to evolve independently twice. 4 Hypothesis 2 is chosen because it requires that each character evolve only once.

FIGURE 23-11 Biologists use the principle of parsimony to choose between alternative hypotheses of evolutionary relationships

ogram with the fewest changes in characters (the one with the fewest homoplasies) be accepted as most probable (FIG. 23-11). In actual practice it is often possible to generate several cladograms that are equally parsimonious. The excluded choices are those with more homoplasies because shared derived characters are more likely than homoplasies. Systematists also use maximum likelihood to make decisions, especially when analyzing molecular data. Maximum likelihood is a statistical method that depends on probability, for example, the probability that nucleotide sequences in DNA and RNA change at a constant rate over time. Complex computer programs analyze large data sets and report the probability of a particular tree.

■ ■

Many evolutionary questions remain. New hypotheses about how organisms are related suggest new taxonomic schemes. Systematics offers tools to help us test these hypotheses. In Chapters 24 through 32, we explore many evolutionary puzzles and discuss current hypotheses about relationships and classifications of organisms.

Review ■ ■ ■

In what way do cladists use shared derived characters in their work? What is outgroup analysis? How do systematists apply the principle of parsimony?

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

23.1 (page 482) 1 State two justifications for the use of scientific names and classifications of organisms.

■

Systematics is the scientific study of the diversity of organisms and their evolutionary relationships. Taxonomy is the branch of systematics devoted to naming, describing, and classifying organisms.

■

■

Classification is the process of arranging organisms into groups based on similarities that reflect evolutionary relationships. Scientific names allow biologists from different countries with different languages to communicate about organisms. Biologists in distant locations must know with certainty that they are studying the same (or different) organisms. Classifications help biologists organize their knowledge.

2 Describe the binomial system of naming organisms and arrange the Linnaean categories in hierarchical fashion from most inclusive to least inclusive. ■ In the binomial system of nomenclature, the basic unit of classification is the species. ■ The name of each species has two parts: the genus name followed by the specific epithet. ■ The hierarchical system of classification includes domain, kingdom, phylum, class, order, family, genus, and species. Each formal grouping at any given level is a taxon.

23.2 (page 483) 3 Describe the three domains and argue for and against classifying organisms in the domains. (Compare this approach with using kingdoms as the main rank of classification.) ■ The three-domain classification system assigns organisms to domains Archaea, Bacteria, or Eukarya. The domain classification is based on molecular data. ■ Domain Eukarya includes the fungi, plants, animals, and protists. ■ Some systematists recognize the kingdoms Archaea, Bacteria, Fungi, Plantae, and Animalia. Several “supergroups” composed of mainly unicellular, mainly aquatic eukaryotic organisms were formerly classified as kingdom Protista. Members of Archaea and Bacteria are prokaryotes. The fungi, which include molds, yeasts, and mushrooms, absorb nutrients produced by other organisms. Kingdoms Plantae and Animalia both consist of multicellular eukaryotes. For additional characteristics of the kingdoms, refer to Table 23-2. As new data are interpreted, organisms assigned to these kingdoms often must be reclassified.

taxa shared a recent common ancestor and how recently they shared that ancestor compared to other groups.

23.3 (page 488) 5 Critically review the difficulties encountered in choosing taxonomic criteria. ■ Systematists seek to determine evolutionary relationships, or phylogeny, based on shared characteristics. Homology, the presence in two or more species of a trait derived from a recent common ancestor, implies evolution from a common ancestor. ■ Some seemingly homologous characters are acquired independently by convergent evolution, independent evolution of similar structures in distantly related organisms, or by reversal, reversion of a trait to its ancestral state. The term homoplasy refers to superficially similar characters that are not homologous. 6 Apply the concept of shared derived characters to the classification of organisms. ■ Shared ancestral characters (plesiomorphies) suggest a distant common ancestor. ■ Shared derived characters (synapomorphies) indicate a more recent common ancestor and can be used as taxonomic criteria. 7 Describe how analyses of molecular homologies contribute to the science of systematics. ■ Molecular systematics depends on molecular structure to clarify phylogeny. Comparisons of nucleotide sequences in DNA and RNA, and of amino acid sequences in proteins, provide important information about how closely organisms are related 8 Contrast monophyletic, paraphyletic, and polyphyletic taxa. 1

2

3

4

5

C

6

E Common D ancestor of groups 4, 5, and 6

Explore the tree of life by clicking on the figures

B

in CengageNOW. A

4 Interpret a cladogram, describing the meaning of its specific nodes and branches.

■

■

■

A monophyletic group, or clade, includes all the descendants of the most recent common ancestor. A paraphyletic group consists of a common ancestor and some, but not all, of its descendants. A polyphyletic group consists of organisms that evolved from different recent ancestors.

23.4 (page 493) 9 Contrast the classification of reptiles and birds by a cladist and by an

■

■

In a cladogram, each branch represents a clade, a group of organisms with a common ancestor. Each node, or branching point, represents the splitting of two or more new groups from a common ancestor. The node represents the most recent common ancestor of the clade represented by the branches. The root represents the most recent common ancestor of all the clades shown in the tree. We can determine the relationships among taxa by tracing along the branches back to the nodes. The cladogram indicates which

evolutionary systematist. ■ Scientists applying the principles of cladistics insist that taxa be monophyletic. Each monophyletic group consists of a common ancestor and all its descendants. Cladists use shared derived characters to determine these relationships. Cladists classify reptiles and birds in a single clade. ■ Biologists using an evolutionary systematics approach classify reptiles and birds in separate classes even though these classes are paraphyletic groups. Evolutionary systematics is based on shared ancestral characters as well as shared derived characters. 10 Describe the construction of a cladogram by using outgroup analysis. ■ Cladists use shared derived characters to reconstruct evolutionary relationships and diagram these relationships in cladograms. They

use outgroup analysis to determine which characters in a given group of taxa are ancestral and which are derived. ■ An outgroup is a taxon that diverged earlier than any of the other taxa being investigated. 11 Apply the principle of parsimony to cladogram construction.

■

In using the principle of parsimony, cladists choose the simplest explanation to interpret the data. Learn more about interpreting and constructing cladograms by clicking on the figures in CengageNOW.

T E S T YO U R U N D E R S TA N D I N G 1. The science of describing, naming, and classifying organisms is (a) systematics (b) taxonomy (c) cladistics (phylogenetic systematics) (d) phenetics (e) evolutionary systematics 2. The mold that produces penicillin is Penicillium notatum. Penicillium is the name of its (a) genus (b) order (c) family (d) species (e) specific epithet 3. Decomposers such as molds and mushrooms belong to which domain and kingdom? (a) Eukarya, Plantae (b) Bacteria, Archaea (c) Archaea, Fungi (d) Eukarya, Archaea (e) Eukarya, Fungi 4. Each branching point in a cladogram (a) is called a root (b) represents a clade (c) represents the divergence of two or more groups from a common ancestor (d) represents horizontal gene transfer (e) marks the divergence of two kingdoms 5. The presence of homologous structures in two different groups of organisms suggests that (a) the organisms evolved from a common ancestor (b) convergent evolution has occurred (c) they belong to a polyphyletic group (d) homoplasy has occurred (e) independently acquired characters may evolve when organisms inhabit similar environments 6. The dolphin and the human both have the ability to nurse their young, whereas the less closely related fish does not. The ability to nurse their young is a (a) shared derived character of mammals (b) shared ancestral character of all vertebrates (c) shared derived character of fish and mammals (d) homologous behavior (e) homoplasy 7. Using specific regions of DNA as a molecular barcode (a) is effective only with bacteria (b) is possible because of similarities in sugars (c) could cause major mistakes in distinguishing among closely related species (d) is an effective way to identify vertical

gene transfer (e) could help taxonomists identify and describe new species 8. A group of organisms that includes a recent common ancestor and all its descendants is (a) polyphyletic (b) paraphyletic (c) monophyletic (d) an example of horizontal gene transfer (e) a sister group 9. Some systematists classify crocodiles and birds in the same taxon because they are contained within a monophyletic group. These systematists follow which approach? (a) phyletic (b) cladistic (c) evolutionary systematic (d) polyphyletic (e) both cladistic and evolutionary systematic 10. In cladistic analysis (a) ancestral characters are used to reconstruct phylogenies (b) characters must be homoplastic (c) polyphyletic groups are preferred (d) the method called maximum likelihood is rarely used (e) outgroup analysis is used to determine a taxon that diverged earlier than the other taxa being studied 11. In interpreting a cladogram (a) we can identify the ancestor of each group by tracing each branch back to the node closest to the root (b) we can identify the taxa to which each group gave rise (c) we must study the relative placement of smaller branches along the main axis of the tree (d) we can determine the relationships by tracing along the branches back to the most recent common ancestor (e) we must identify horizontal gene transfer 12. When cladists apply the principle of parsimony, they (a) choose the cladogram that requires the fewest changes in characters as the most probable explanation to interpret the data (b) select multiple hypotheses to explain each relationship (c) do not use outgroups (d) typically use a polyphyletic approach (e) hypothesize that the most complex explanation is most probably the correct one

CRITICAL THINKING 1. What are the advantages of PhyloCode? of barcoding organisms? 2. What are some of the criteria/tools biologists might use to assign the protists to monophyletic groups? 3. EVOLUTION LINK. Explain why evolution is not always linear. 4. EVOLUTION LINK. Are members of a clade similar because they share a common ancestor, or do they belong to the same clade because they are similar? How might your answer vary depending on which approach to systematics you are following? 5. EVOLUTION LINK. The TATA-binding protein (TBP) is thought to be necessary for transcription in all eukaryotic cell nuclei. Studies show that archaea, but not bacteria, have a pro-

tein structurally and functionally similar to TBP. What does this similarity suggest regarding the evolution of archaea, bacteria, and eukaryotes? How might knowledge of this similarity affect how systematists classify these organisms? 6. ANALYZING DATA. Construct a cladogram based on the following data. Mosses are plants with no vascular tissue. Horsetails, ferns, gymnosperms (pines and other plants with naked seeds), and angiosperms (flowering plants) are all vascular plants. Seeds are absent in all but the gymnosperms and angiosperms. Angiosperms are the only seed plants with flowers. (Hint: To help you construct the cladogram, draw a simple table showing which characters are present in each group. See Figure 23-9.)

7. ANALYZING DATA. In the illustration, what kind of grouping is represented by the bracketed area? What type of group is formed by 2 and 3? What type of group is formed by 2, 4, and 6? Taxon 1

2

3

4

5

6

8. SCIENCE, TECHNOLOGY, AND SOCIETY. A nurse and two patients in the same hospital die from an unknown bacterial infection that does not respond to penicillin. How would you go about determining the phylogeny of this bacterium? (Consider strategies learned in this chapter.) How might knowing its phylogeny be helpful in this situation? Additional questions are available in CengageNOW at www.cengage.com/ login.

24

Viruses and Subviral Agents

NIBSC/Science Photo Library/Photo Researchers, Inc.

This colorized SEM shows human immunodeficiency virus (HIV; red) infecting a T helper cell (an immune system cell; green).

KEY CONCEPTS

D

uring the late 1800s, botanists searched for the cause of tobacco mosaic disease, which stunts the growth of tobacco plants and gives

24.1 A virus is a small particle consisting of a nucleic acid

the infected tobacco leaves a spotted, mosaic appearance. They found

core surrounded by a protein coat; in order to reproduce, a virus must infect a living cell.

they could transmit this disease to healthy plants by daubing their leaves

24.2 Viruses can be classified based on their host range,

botanist, showed that the sap was still infective after it had been passed

what type of nucleic acid they have, and whether the nucleic acid is single-stranded or double-stranded.

24.3 In a lytic reproductive cycle, a virus uses the host cell’s molecular machinery to replicate itself, destroying the host cell in the process; in a lysogenic cycle, the viral genome is integrated into the host DNA and is then called a “provirus.”

24.4 Viruses infect the cells of all kinds of organisms; they cause serious diseases in plants and animals.

24.5 According to the cellular origin hypothesis, viruses

with the sap of diseased plants. In 1892, Dmitri Ivanowsky, a Russian through porcelain filters designed to filter out all known bacteria. A few years later, in 1898, Martinus Beijerinck, a Dutch microbiologist, provided evidence that the agent that caused tobacco mosaic disease had many characteristics of a living organism. However, it appeared that the infective agent could reproduce only within a living cell. Beijerinck named the infective agent, virus (from the Latin word virus, which means “poison”). Tobacco mosaic virus was the first virus to be recognized. Early in the 20th century, scientists discovered other infective agents,

may have originated as mobile genetic elements such as transposons or plasmids.

like those responsible for tobacco mosaic disease, that could cause dis-

24.6 Subviral agents, which include satellites, viroids, and

could not be seen with the light microscope. They also passed through fil-

prions, are smaller and simpler than viruses.

ters that removed all known bacteria. Curiously, they could not be grown

ease in animals or kill bacteria. These pathogens were so small that they

in laboratory cultures unless living cells were present. The development of the electron microscope in the 1930s made it possible to see viruses for the first time. Most of the viruses that infect animals, plants, and bacteria were identified during the second half of the 20th century. Viruses cause many plant diseases and are responsible for billions of dollars in crop losses each year. You may be familiar with some of the many human viral diseases, including influenza. smallpox, rubella (German measles), mumps, rabies, polio, herpes, and acquired human immunodeficiency syndrome (AIDS; see figure).

Viruses are very small As early investigators reported, viruses are indeed very small. Most range in size from 20 to 300 nm. (A nanometer is one-thousandth of a micrometer.) The poliovirus is about 30 nm in diameter (about the size of a ribosome). If we could line these viruses up end to end, it would take almost a million of them to span one inch! A larger virus, such as the poxvirus that causes smallpox, can measure up to 300 nm long and 200 nm wide. The largest known virus is Mimivirus, with a diameter of about 400 nm, the size of a small bacterium (mycoplasma). Mimivirus, which was discovered in 2003, infects amoebas.

Viruses are the most numerous microorganisms on Earth, and they influence many ecological processes. For example, viruses kill large amounts of marine biomass each day, contributing significantly to the recycling of nutrients. Viruses adapt to their environments through evolution. They also influence the biodiversity of other organisms. Viruses can transfer their own genes into the genetic material of their hosts. They can also transfer eukaryotic genes from one organism to another, crossing species boundaries. In this chapter we examine the characteristics and diversity of viruses, and the smaller subviral agents, viroids and prions. We include viruses in the diversity unit of Biology because they have some characteristics of living things and because they impact all living organisms.

24.1 THE STATUS AND STRUCTURE OF VIRUSES ■ ■ LEARNING OBJECTIVES 1 2

Contrast a virus with a cellular organism. Describe the structure of a virus.

A virus is a very small infective agent that consists of a core of nucleic acid and is dependent on a living host. The study of viruses is called virology, and biologists who study viruses are virologists. Most biologists view viruses as nonliving particles because they are not composed of cells and they cannot carry on metabolic activities or reproduce on their own. They do not have the components necessary to carry on cellular respiration or to synthesize proteins and other molecules. Viruses do contain the nucleic acids necessary to make copies of themselves. They replicate by invading living cells and commandeering their metabolic machinery. In order to multiply, a virus must infect a cell in which it can replicate. Thus, viruses are obligate intracellular parasites; they survive only by using the resources of a host cell, the cell the virus invades. Viruses infect all types of organisms, including bacteria, archaea, protists, plants, fungi, and animals. Some viruses even infect other viruses. Interestingly, viruses evolve by natural selection. For these reasons, most biologists view viruses as being at the edge of life, although a few argue that they are very simple life-forms.

A virus consists of nucleic acid surrounded by a protein coat The nucleic acid core of the virus is surrounded by a protein coat called a capsid. Microbiologists use the term virion to refer to the complete virus particle that is outside a host cell. The virion is the form in which the virus moves from the cell in which it was produced to a new host cell in which it can replicate its genome. However, here we will use the term virus in a broad sense, even when referring to virions. A typical virus contains either deoxyribonucleic acid (DNA) or ribonucleic acid (RNA), not both. The nucleic acid can be single-stranded or double-stranded. Thus, a virus can have singlestranded (ss) DNA, double-stranded (ds) DNA, ssRNA, or dsRNA. As we will discuss, the type of nucleic acid is important in classifying viruses. The virus genome typically consists of 5000 to more than 100,000 bases or base pairs (depending on whether it is singlestranded or double-stranded). Again, Mimivirus is an exception. Investigators have sequenced the genome of Mimivirus and reported that this large double-stranded DNA virus has more than one million base pairs. Mimivirus also has six transfer RNAs.

The capsid is a protective protein coat The capsid consists of protein subunits, called capsomers. The capsomers determine the shape of the virus. Capsids generally are either helical or polyhedral, or a combination of both shapes. Helical viruses, such as the tobacco mosaic virus (TMV), appear as long rods or threads (FIG. 24-1a). Its capsid is a hollow cylinder made up of proteins that form a groove into which the RNA fits. Polyhedral viruses, such as the adenoviruses (which cause a number of human illnesses, including some respiratory infections), appear somewhat spherical (FIG. 24-1b). Its capsomers are organized in equilateral triangles. The capsid of a large virus can consist of several hundred capsomers. The most common polyhedral structure is an icosahedron, a structure with 20 identical surface faces (each face is a triangle). The human immunodeficiency virus (HIV) that causes acquired immunodeficiency syndrome (AIDS) is an enveloped virus. Proteins in the envelope extend outward like spikes (FIG. 24-1c). Some viruses have both helical and polyhedral components. Viruses that infect bacteria are called bacteriophages, or simply phages (FIG. 24-1d). The T4 phage that infects the bacterium Esch-

Biozentrum/Science Photo Library/ Photo Researchers, Inc.

© Dennis Kunkel Microscopy, Inc. / Visuals Unlimited / Corbis

RNA inside capsid

(b) Color-enhanced TEM of an adenovirus. The capsid is composed of 252 subunits (visible as tiny ovals) arranged into a 20-sided polyhedron. Twelve of the subunits have projecting protein spikes that permit the virus to recognize host cells.

Capsomers

© Dr. Klaus Boller / Photo Researchers, Inc.

Enzymes

Capsid RNA

© Department of Microbiology, Biozentrum, University of Basel / Photo Researchers, Inc.

200 nm

Envelope Envelope protein

DNA inside capsid

50 nm

Capsid

(a) TEM of tobacco mosaic virus. This virus is rod-shaped with a helical arrangement of capsid proteins.

Capsid with antenna-like protein spikes

DNA inside capsid

Tail

Tail fibers Emerging DNA

100 nm

50 nm

(c) Color-enhanced TEM of HIV, the virus that causes AIDS. The virus is leaving a host cell (pink). The virus is enclosed in an envelope (green) derived from the host’s plasma membrane. Viral proteins project from the envelope. The capsid of the virus is shown in yellow.

Capsid

(d) Color-enhanced TEM of T4 bacteriophage. This virus has a polyhedral head and a helical tail. The virus attaches to the cell wall of the bacterial host by its tail fibers.

FIGURE 24-1 Animated The structure of a virus A virus consists of a DNA or RNA core surrounded by a protein coat called a capsid. The capsid is made up of protein subunits called capsomers. Some viruses have an outer membranous envelope surrounding the capsid.

erichia coli consists of a polyhedral “head” attached to a helical “tail.” Many phages have this shape. Many phages also have tail fibers that attach to the host cell.

Some viruses are surrounded by an envelope Some viruses, called enveloped viruses, have an outer membranous envelope that surrounds the capsid. Typically, the virus acquires the envelope from the host cell’s plasma membrane as it leaves the host cell (FIG. 24-2). Interestingly, while inside the host cell, the virus synthesizes certain proteins and inserts them into the host’s

plasma membrane. Thus, the viral envelope consists of phospholipids and proteins of the host’s plasma membrane, as well as distinctive proteins produced by the virus itself. Some viruses produce envelope proteins that extend out from the envelope as spikes. As we will discuss, these spikes can be very important in the virus’s interaction with the host cell.

Review ■ ■

What characteristics of life are absent in a virus? What are the structural components of a virus?

Host cell

Nucleic acid Capsomers

(a) Naked virus Plasma membrane of host cell

Envelope Protein spikes

Glycoproteins produced by virus

(b) Enveloped virus

FIGURE 24-2 Comparison of naked and enveloped viruses As the virus leaves the host cell, the host’s plasma membrane wraps around the virus, forming the envelope. The envelope contains proteins produced by the virus.

24.2 CLASSIFICATION OF VIRUSES ■ ■ LEARNING OBJECTIVE 3

Identify three characteristics used to classify viruses.

Viruses present a taxonomic challenge to biologists because they do not have the characteristics that define living organisms (see Chapter 1). They are acellular, do not carry on metabolic activities, and reproduce only by taking over the reproductive machinery of other cells. Viruses do not manufacture proteins and so do not have distinctive rRNA. For these reasons, viruses are not classified in any of the three domains. Viruses can be classified based on their host range, the types of organisms they infect. They may be referred to as plant viruses, animal viruses, bacterial viruses, and so on. Viruses are more formally classified into taxa from species to orders. The International Committee on Taxonomy of Viruses (ICTV), a group of virologists, decides on specific criteria for classifying and naming viruses. Recently, based on host range and other characteristics, the ICTV classified viruses into 5 orders, 84 families, 307 genera, and more than 2000 species. Virus family names include the suffi x -viridae. Note that this classification system is not a tra-

ditional Linnaean system; it does not assign viruses to domains, kingdoms, or phyla. The Baltimore classification system classifies viruses based on the type of nucleic acid the virus contains, whether the nucleic acid is single-stranded or double-stranded, and on how mRNA is produced. Other traits considered in viral classification are the size and shape of the virus, the presence of an envelope, and the method by which the virus is transmitted from host to host.

Review ■

What are three characteristics used to classify viruses?

24.3 VIRAL REPLICATION ■ ■ LEARNING OBJECTIVES 4 Characterize bacteriophages. 5 Contrast a lytic cycle with a lysogenic cycle.

As we have discussed, viruses reproduce, but only within the complex environment of the living host cells they infect. Viruses use their genetic information to force their host cells to replicate their viral nucleic acid and to make the proteins they need. They

take over the transcriptional and translational mechanisms of the host cell.

KEY POINT

Viruses reproduce by seizing control of the metabolic machinery of a host cell. The host cell is destroyed in a lytic infection.

Bacteriophages infect bacteria Phages

Much of our knowledge of viruses has come from studying bacteriophages (“bacteria eaters”), more simply referred to as phages. These viruses can be cultured easily within living bacteria in the laboratory. Microbiologists have identified more than 2000 phages. Bacteriophages are among the most complex viruses (see Fig. 24-1d). Their most common structure consists of a long nucleic acid molecule (usually dsDNA) coiled within a polyhedral head. Most have a tail, which may be contractile and may function in penetration of the host cell. Before the age of sulfa drugs and antibiotics, phages were used clinically to treat infection. In the 1940s, they were abandoned (at least in Western countries) in favor of antibiotics, which were more dependable and easier to use. Now, with the widespread and growing problem of bacterial resistance to antibiotics, these bacteria-killing viruses are once again the focus of research. With new knowledge of phages and sophisticated technology, several research groups are investigating phages to determine which ones kill which species of bacteria. Scientists are genetically engineering phages so that bacteria will be slower to evolve resistance to them. Phages can also be used to improve food safety. For example, certain phages can kill deadly strains of E. coli in cattle. These bacteria do not appear to make cattle ill but can cause illness and death in people who eat contaminated, undercooked hamburgers.

1

Bacterium

Bacterial DNA 2

3

Replication and synthesis. Phage DNA is replicated. Phage proteins are synthesized.

4

Assembly. Phage components are assembled into new viruses.

5

Release. Bacterial cell lyses and releases many phages that can then infect other cells.

1. Attachment (or adsorption). The virus attaches to specific receptors on the host cell. This process ensures that the virus infects only its specific host. 2. Penetration. The virus penetrates the host plasma membrane and moves into the cytoplasm. Many viruses that infect animal cells enter the host cell intact. Some phages inject only their nucleic acid into the cytoplasm of the host cell; the capsid remains on the outside.

(a) The sequence of events in a lytic infection.

© Oliver Meckes/MPL-Tübingen / Photo Researchers, Inc.

Lytic reproductive cycles destroy host cells In a lytic cycle, the virus lyses (destroys) the host cell. When the virus infects a susceptible host cell, it forces the host to use its metabolic machinery to replicate viral particles. Viruses that have only a lytic cycle are described as virulent, which means that they cause disease, and often death. Five steps are typical in lytic viral reproduction (FIG. 24-3):

Penetration. Phage DNA enters bacterial cell.

Phage protein Phage DNA

Viruses replicate inside host cells Viruses infect bacterial, plant, and animal cells in basically similar ways. We will focus on phage infection of bacteria because this process is best understood. The reproductive cycle of viruses begins with a virus coming into contact with a host cell. The virus typically attaches to the surface of the host cell. The viral nucleic acid must enter the host cell and synthesize the components it needs to reproduce itself. Then, viral components are assembled and viruses are released from the cell—ready to invade other cells. Two types of viral reproductive cycles are lytic and lysogenic cycles.

Attachment. Phage attaches to cell surface of bacterium.

80 nm

(b) Color-enhanced TEM of phages infecting a bacterium, Escherichia coli.

FIGURE 24-3 Animated Lytic cycle In a lytic cycle, the virus destroys the host cell.

3. Replication and synthesis. The viral genome contains all the information necessary to produce new viruses. Once inside, the virus degrades the host-cell nucleic acid and uses the molecular machinery of the host cell to replicate its own nucleic acid and to produce viral proteins. Many antiviral drugs interfere with replication of viral nucleic acid. 4. Assembly. The newly synthesized viral components are assembled into new viruses. 5. Release. Assembled viruses are released from the cell. Generally, lytic enzymes, produced by the phage late in the replication process, destroy the host plasma membrane. Phage release typically occurs all at once and results in rapid cell lysis. In contrast, animal viruses are often released slowly or bud off from the plasma membrane.

KEY POINT

1

2

Once released, the viruses infect other cells, and the process begins anew. The time required for viral reproduction, from attachment to the release of new viruses, varies from less than 20 minutes to more than an hour. How do bacteria protect themselves from phage infection? You may recall from Chapter 15 that bacteria produce restriction enzymes, enzymes that cut up the foreign DNA of the phage; this prevents the phage DNA from duplicating. The bacterial cell protects its own DNA by slightly modifying it after replication so that the restriction enzyme does not recognize the sites it would cut. In lysogenic cycles, temperate viruses integrate into the host DNA Temperate viruses do not always destroy their hosts. In a lysogenic cycle, the viral genome usually becomes integrated into the host bacterial DNA. The integrated virus is called a prophage or provirus. When the bacterial DNA replicates, the prophage also replicates (FIG. 24-4). The viral genes that code for viral structural proteins may be repressed indefinitely. Bacterial cells carrying prophages are called lysogenic cells. Certain external conditions (such as ultraviolet light and X-rays) cause temperate viruses to revert to a lytic cycle and then destroy their host. Sometimes temperate viruses become lytic spontaneously. Bacterial cells containing certain temperate viruses may exhibit new properties. This change is called lysogenic conversion. An interesting example involves the bacterium Corynebacterium diphtheriae, which causes diphtheria. Two strains of this species exist, one that produces a toxin (and causes diphtheria) and one that does not. The only difference between these two strains is that the toxin-producing bacteria are infected by a specific temperate phage. The phage DNA codes for the powerful toxin that causes the symptoms of diphtheria. Similarly, the bacterium Clostridium botulinum, which causes botulism, a serious form of food poisoning, is harmless unless it contains certain prophage DNA that induces synthesis of the toxin.

Review ■

■

What are the steps in a lytic cycle? Draw diagrams to illustrate your answer. How is a lysogenic cycle different from a lytic cycle?

In a lysogenic infection, temperate phages integrate their nucleic acid into the host genome without destroying the host cell.

Attachment. Phage attaches to cell surface of bacterium.

Penetration. Phage DNA enters bacterial cell.

Prophage 3

Integration. Phage DNA integrates into bacterial DNA.

4

These cells may exhibit new properties.

Replication. Integrated prophage replicates when bacterial DNA replicates.

FIGURE 24-4 Animated Lysogenic cycle Temperate phages integrate their nucleic acid into the host-cell DNA, which then becomes a lysogenic cell.

24.4 VIRAL DISEASES ■ ■ LEARNING OBJECTIVES 6 Contrast viral infection of plants and animals, and identify specific 7

diseases caused by viruses. Describe the reproductive cycle of a retrovirus, such as human immunodeficiency virus (HIV).

Sir Peter Medawar, 1960 Nobel laureate in Medicine, defined a virus as “a piece of bad news wrapped in protein.” Viruses that are pathogens are responsible for some very serious diseases, including smallpox, rabies, influenza (flu), and AIDS. However, although viruses must infect cells in order to reproduce, most known viruses do not cause disease. Here we focus on viral infection of plants and animals.

Viruses cause serious plant diseases Viruses cause many important plant diseases and are responsible for billions of dollars in agricultural losses and lower crop quality

© Jack M. Bostrack/Visuals Unlimited, Inc.

Kenneth M. Corbett

worldwide each year. Infected crops almost always produce lower yields. Viral infections do not usually kill plants, but they do stunt their growth, cause changes in the shape of the foliage, and may cause spots, streaks, or mottled patterns on leaves, flowers, or fruits (FIG. 24-5). Most plant viruses have capsids, but do not have envelopes. The genome of most plant viruses consists of single-stranded RNA. Plant viruses are typically named according to the type of plant they infect and by their effects on the plant. Tobacco mosaic virus was the first virus ever discovered. Insects are important vectors (b) Pepper leaves infected with tobacco (a) Virus-streaked tulip. The virus that causes of plant disease. As they feed on mosaic virus. The leaf is characteristically this relatively harmless disease affects pigment plant tissues, aphids, leaf-hoppers, mottled with light green areas. formation in the petals. and many other insects spread viral diseases among plants. Because FIGURE 24-5 Plant diseases caused by viruses of plants’ thick cell walls, viruses cannot penetrate plant cells unless ease. Living conditions, including sanitation, nutrition, physical the cells are damaged. Plant viruses can also be transmitted from stress, level of health care, and sexual practices, are important facone generation to the next through infected seeds or by asexual tors in the spread of disease. In the United States and other highly propagation. Once a plant is infected, the virus spreads through developed countries, infectious disease accounts for about 4% to the plant body by passing through plasmodesmata (cytoplasmic 8% of deaths compared with death rates of 30% to 50% in develconnections) that penetrate the walls between adjacent cells (see oping regions. Even at the level of our current knowledge about Fig. 5-27). viruses and epidemiology, just how prepared are we to contain a There are no known cures for most viral diseases of plants, so particularly virulent virus? How well are we containing HIV (the infected plants are commonly burned. Many agricultural scientists virus that causes AIDS), which continues to infect and kill millions are focusing their efforts on preventing viral disease by developing of people worldwide every year? virus-resistant strains of important crop plants. Since the September 11, 2001, terrorist attacks in the United States, bioterrorism has become a critical concern worldwide. BioViruses cause serious diseases in animals terrorism is the intentional use of microorganisms or toxins deHundreds of different viruses infect animals, including humans. rived from living organisms to cause death or disease in humans, Animal viruses cause hog cholera, foot-and-mouth disease, canine animals, or plants on which humans depend. Terrorists could condistemper, influenza, and certain types of cancer (such as feline ceivably initiate epidemics of smallpox, anthrax, plague, and other leukemia and cervical cancer). Viruses cause chickenpox, herpes potentially fatal diseases. (Note that anthrax and plague are caused simplex (one type causes genital herpes), mumps, rubella (Gerby bacteria; see Table 25-5 in Chapter 25.) The quest for effecman measles), rubeola (measles), rabies, warts, infectious monotive treatments and vaccines for these diseases has taken on new nucleosis, influenza, viral hepatitis, and AIDS (TABLE 24-1 and urgency. Inquiring About: Influenza and Other Emerging Diseases). Most huMany animal viruses have lytic reproductive cycles mans suffer from two to six viral infections each year, including Most viruses cannot survive very long outside a living host cell, common colds. so their survival depends on their being transmitted from one aniAs discussed in Chapter 1, the outbreak of H1N1 influenza in mal to another. However, their host range may be quite limited 2009 was met by rapid responses from the World Health Organibecause attachment to a host cell is very specific. The type of atzation (WHO), U.S. Centers for Disease Control and Prevention tachment proteins on the surface of a virus determines what type (CDC), and other public health agencies throughout the world. of cell it can infect. Receptors typically vary with each species and Careful surveillance and willingness of countries to rapidly share sometimes with each type of tissue. Thus, many human viruses information helped minimize the spread of this pandemic. Effeccan infect only humans because the viral attachment proteins tive vaccines and treatment protocols were quickly developed. combine only with receptor sites found on human cell surfaces. Human activity, including social factors such as urbanization, The measles virus and poxviruses infect many types of human global travel, and war, contributes to epidemics of infectious dis-

TABLE 24-1

Some Viruses That Infect Vertebrates

Group

Diseases Caused

Characteristics

DNA Viruses with Envelope Poxviruses Smallpox, cowpox,* monkeypox, and economically important diseases of domestic fowl Herpesviruses Cold sores (herpes simplex virus type 1); genital herpes, a sexually transmitted disease (herpes simplex virus type 2); chickenpox and shingles (herpes varicella–zoster virus); infectious mononucleosis and Burkitt’s lymphoma (Epstein–Barr virus) DNA Viruses with No Envelope Adenoviruses Respiratory tract disorders (e.g., sore throat, tonsillitis), conjunctivitis, and gastrointestinal disorders are caused by more than 40 types of adenoviruses in humans; other varieties infect other animals Papovaviruses‡ Human warts and some degenerative brain diseases; some cancers, including cervical cancer Parvoviruses Infections in dogs, swine, arthropods, rodents; gastroenteritis in humans (transmitted by consumption of infected shellfish) RNA Viruses with Envelope Togaviruses Rubella (German measles)

Orthomyxoviruses

Influenza (flu) in humans and other animals

Paramyxoviruses Rhabdoviruses Coronaviruses Flaviviruses

Rubeola (measles) and mumps in humans; distemper in dogs Rabies Upper respiratory infections; SARS Yellow fever; West Nile virus; hepatitis C (the most common reason for liver transplants in the United States) Hemorrhagic fever, including that caused by the Ebola virus St. Louis encephalitis; hantavirus pulmonary syndrome (caused by Sin Nombre virus, a hantavirus) AIDS; some types of cancer

Filoviruses Bunyaviruses Retroviruses

RNA Viruses with No Envelope Picornaviruses Polio (poliovirus); hepatitis A (hepatitis A virus); intestinal disorders (enteroviruses); common cold (rhinoviruses); aseptic meningitis (coxsackievirus, echovirus) Reoviruses Vomiting and diarrhea; encephalitis

dsDNA; large, complex viruses; replicate in the cytoplasm of the host cell dsDNA; medium to large, enveloped viruses; replicate in the host nucleus†

dsDNA; replicate in the host nucleus

dsDNA ssDNA; some require a helper virus in order to multiply

ssRNA that can serve as mRNA; large diverse group of medium-sized, enveloped viruses; many transmitted by arthropods ssRNA that serves as template for mRNA synthesis; medium-sized viruses that often exhibit projecting spikes ssRNA; resemble orthomyxoviruses but somewhat larger ssRNA ssRNA; largest known RNA virus ssRNA ssRNA ssRNA ssRNA viruses that contain reverse transcriptase for transcribing the RNA genome into DNA; two identical molecules of ssRNA

ssRNA that can serve as mRNA; diverse group of small viruses dsRNA

*The vaccinia (cowpox) virus is used to produce genetically engineered vaccines. †

These viruses frequently cause latent infections; some cause tumors.

‡

The virus SV40 has been used as a vector to transport genes into cells.

tissue because their attachment proteins combine with receptor sites on a variety of cells. In contrast, the poliovirus attaches to specific types of human cells, such as those that line the digestive tract and motor neurons of the brain and spinal cord. Some viruses, such as the adenoviruses, have fibers that project from the capsid and adhere to complementary receptors on the host cell. Other viruses, such as those that cause herpes and rabies, are surrounded by a lipoprotein envelope with projecting glycoprotein spikes that attach to a host cell. The influenza virus has rodlike spikes that project from its envelope. The glycoprotein spikes bind with specific receptors present only on cells lining the

respiratory tract of certain vertebrates, including wild and domestic birds, horses, pigs, and humans. Influenza vaccine prevents attachment by stimulating the host’s antibodies to cover the spikes projecting from the virus. Viruses have several ways to penetrate animal cells (FIG. 24-6). After attachment to a host-cell receptor, some enveloped viruses fuse with the animal cell’s plasma membrane. The entire virus, including the capsid and nucleic acid, is released into the animal cell. Other viruses enter the host cell by endocytosis. In this process, the plasma membrane of the animal cell invaginates to form a membrane-enclosed vesicle that contains the virus. Endocytosis is

In q u ir in g A b o u t I N F LU E N Z A A N D OT H E R E M E R G I N G D I S E A S E S How do new strains of the influenza virus re-emerge? Emerging diseases, those new to the human population—such as AIDS, severe acute respiratory syndrome (SARS), Ebola, eastern equine encephalitis, and West Nile virus—often appear suddenly. Many new, continual, or re-emerging pathogens have the potential to strike globally. Re-emerging diseases are those that have been almost eradicated and then strike unpredictably, causing an epidemic, sometimes in a new geographic area. Many familiar diseases, such as influenza, malaria, tuberculosis, and bacterial pneumonias, continue to infect large numbers of people and increasingly reappear in forms that are resistant to drug treatments. Drug-resistant pathogens are a major challenge to public health and other health-care professionals. Historically, new viral strains have claimed many human lives. For example, in 1918 an influenza (flu) pandemic killed more than 20 million people throughout the world. Flu pandemics also occurred in 1957 and 1968. Each year new strains of influenza virus evolve and become infectious. If the new combination of viral genes is unfamiliar to the human immune system, it may spread easily, resulting in a pandemic. According to the U.S. Centers for Disease Control and Prevention (CDC), influenza kills about 36,000 individuals in the United States every year. New strains of influenza are thought to infect one species, such as a species of birds. At first, they are not able to spread among mammals. However, a virus of the avian strain can mutate and become virulent. The avian virus can also exchange RNA with a virus that has the genetic material necessary to spread among swine or other mammals. Swine can be susceptible to both avian and human strains of the virus. Inside a swine’s body, the viruses can exchange bits of RNA and develop the ability to infect humans. Such reassortment can also occur inside a human host and produce a viral strain that can spread from person to person (see figure). H1N1 influenza evolved in this way.

Influenza virus not infectious to humans

Mutant virus that can infect humans Mutation

1

2

Virus can infect human cells but cannot spread to other humans.

Influenza virus may mutate to forms that can infect human cells.

Viral RNA

Influenza viruses from two different strains infect human cell by releasing their RNA into cell. Neither of these viruses can spread from human to human.

3

Nucleus

RNA from both viruses is duplicated in nucleus (duplicated RNA strands not shown).

Host cell

4

New viruses assembled with reassorted RNA; new viruses have RNA from both strains.

Virus with reassorted RNA 5

Viruses of new strain leave host cell and now are highly infectious to humans.

An influenza virus such as H1N1 evolves as it mutates and its RNA reassorts in the bodies of birds, swine, and humans. Eventually, it becomes virulent.

advantageous to the virus because the endocytotic vesicle delivers the virus deep into the cytosol. In DNA animal viruses, replication of viral DNA and protein synthesis are similar to the processes by which the host cell would normally carry out its own DNA replication and protein synthesis. In most RNA viruses, RNA synthesis takes place with the help of an RNA-dependent RNA polymerase. Retroviruses are RNA viruses that have a DNA polymerase called reverse transcriptase, which transcribes the RNA genome

into a DNA intermediate (FIG. 24-7). This DNA becomes integrated into the host DNA by an enzyme also carried by the virus. Copies of the viral RNA are synthesized as the incorporated DNA is transcribed by host RNA polymerases. The human immunodeficiency virus (HIV) that causes acquired immunodeficiency syndrome (AIDS) is a retrovirus. Certain cancer-causing viruses are also retroviruses. After viral genes are transcribed, the viral structural proteins are synthesized. The capsid is produced, and then assembly of new virus

KEY POINT

Viruses enter animal cells by membrane fusion or by endocytosis. Envelope proteins

1

Envelope Capsid

Virus attaches to specific receptors on plasma membrane of host cell.

Nucleic acid

2

Membrane fusion. Viral envelope fuses with plasma membrane.

3

Virus is released into host-cell cytoplasm.

4

Viral nucleic acid separates from its capsid.

5

Viral nucleic acid enters hostcell nucleus and replicates.

Receptors Host-cell plasma membrane

Capsid

Cytoplasm Nucleus Centers for Disease Control and Prevention, U.S. Department of Health and Human Services

Nucleic acid Ribosomes

ER Viral proteins

(a) This child’s rash is a symptom of rubella (German measles), which is caused by an RNA virus spread by close contact. 10

mRNA

6

7

Viruses are released from host cell.

Viral nucleic acid is transcribed into mRNA.

Host ribosomes are directed by mRNA to synthesize viral proteins.

Glycoproteins 8

9

Vesicles transport glycoproteins to host-cell plasma membrane.

New viruses are assembled and enveloped by host-cell plasma membrane.

(b) Some viruses enter animal cells by membrane fusion. Replication occurs, and the new viruses are released. 3

Host-cell plasma membrane

Endosomal vesicle forms and moves into cytoplasm.

5

Host-cell plasma membrane surrounds virus. 1 Virus makes contact with plasma membrane of host cell. Host-cell cytoplasm 2

(c) Some viruses enter the host cell by endocytosis.

FIGURE 24-6 Some viruses cause diseases in animals

4

Viral envelope fuses with host-cell plasma membrane (not shown).

Virus is released into host-cell cytoplasm.

KEY POINT

Retroviruses use reverse transcriptase to transcribe their RNA into DNA; the viral DNA becomes part of the host DNA. When activated, viral DNA uses host enzymes to transcribe viral RNA. HIV

Envelope protein

Nucleic acid (RNA)

1

HIV attaches to host-cell plasma membrane.

Envelope Capsid Enzymes (reverse transcriptase, ribonuclease, integrase, protease)

2

HIV enters host-cell cytoplasm.

CD4 Receptors Viral nucleic Reverse acid (RNA) transcriptase

Host-cell plasma membrane

3

Cytoplasm ssDNA Nucleus

4

dsDNA

Host chromosome 5

Viral RNA

6

7

8

Capsid is removed by enzymes. Reverse transcriptase catalyzes synthesis of single-stranded (ss) DNA that is complementary to viral RNA.

The ssDNA then serves as template for synthesis of complementary DNA strand, resulting in double-stranded (ds) DNA.

dsDNA is transferred to host nucleus and enzyme integrase integrates DNA into host chromosome.

When activated, viral DNA uses host enzymes to transcribe viral RNA.

Viral RNA leaves nucleus, viral proteins are synthesized on host ribosomes, and virus is assembled.

Virus buds from host cell, using host-cell plasma membrane to make viral envelope.

FIGURE 24-7 Animated Life cycle of HIV, the retrovirus that causes AIDS HIV infects T helper cells, specialized cells of the host’s immune system. The virus attaches to protein receptors, known as CD4, on the plasma membrane of T helper cells. HIV has two identical single-stranded RNA molecules.

particles takes place. Finally, release occurs. Viruses that do not have an outer envelope exit by cell lysis. The plasma membrane ruptures, releasing many new viral particles. Enveloped viruses obtain their lipoprotein envelopes by picking up a fragment of the host plasma membrane as they leave the infected cell (see Figs. 24-6b and 24-7).

Viral proteins damage the host cell in several ways. These proteins may alter the permeability of the plasma membrane or may inhibit synthesis of host nucleic acids or proteins. Viruses sometimes damage or kill their host cells by their sheer numbers. A poliovirus can produce 100,000 new viruses within a single host cell!

Viruses quickly become resistant to antiviral drugs Antibiotics are specific for fighting bacteria; they do not kill viruses. However, researchers have developed several antiviral drugs. Most of these drugs inhibit the replication of many RNA and DNA viruses. Amantadine has been used mainly to treat patients with influenza. The drug inhibits penetration or uncoating of viral nucleic acids. However, recent strains of influenza virus have become resistant to Amantadine, perhaps as a result of using the drug in poultry feed in certain areas of Asia. A newer type of antiviral drug, including Tamiflu, inhibits neuraminidase, a viral enzyme necessary for the virus to leave the host cell. However, viruses are quickly becoming resistant to Tamiflu. In 2008, the Centers for Disease Control and Prevention found that about 11% of individuals with the most common type of influenza had Tamiflu-resistant strains. In 2009, 99% of individuals with this influenza had Tamiflu-resistant strains! It is thought that a single mutation that occurs spontaneously is responsible for this resistance. Other antiviral drugs are in clinical trials. Some inhibit viral attachment to host cells, and others interfere with replication of viral nucleic acid.

Review ■ ■ ■ ■

How do viruses infect plant cells? What are the mechanisms by which viruses enter animal cells? How do viruses damage animal cells? How do retroviruses such as HIV make copies of themselves?

24.5 EVOLUTION OF VIRUSES ■ ■ LEARNING OBJECTIVE 8

Trace the evolutionary origin of viruses according to current hypotheses and describe recent research regarding the evolution of polydnaviruses.

Where did viruses come from? The most widely held hypothesis is that viruses derive from bits of nucleic acid that “escaped” from cellular organisms. According to this cellular origin hypothesis, viruses may have originated as mobile genetic elements such as transposons (see Chapter 13) or plasmids (small, circular DNA fragments discussed in Chapter 15 and Chapter 25). Such fragments could have moved from one cell and entered another through damaged cell membranes. According to this hypothesis, some viruses may trace their origin to animal cells, others to plant cells or to bacterial cells. Their multiple origins may explain why many viruses are species specific; perhaps they infect only those species that are closely related to the organisms from which they originated. This hypothesis is supported by the genetic similarity between some viruses and their host cells—a closer similarity than exists between one type of virus and another. According to the coevolution hypothesis, viruses appeared early in the history of life, even before the three domains diverged. Evidence for this hypothesis comes from similarities found in the protein structures of some viral capsids and in genetic similari-

ties between some viruses that infect archaea and some that infect bacteria. Molecular biologists studying this hypothesis consider it improbable that these similarities evolved independently. This evidence suggests that viruses diverged very early. However, viruses are parasites, dependent on their hosts. How could they have existed before their hosts evolved? According to the regressive hypothesis, viruses evolved from small, independent cells that were parasites in larger cells. Genes that they did not need, like those for protein synthesis, were gradually lost through evolution. This hypothesis is supported by certain bacteria (chlamydia and rickettsia) that are able to reproduce only inside the cells of their hosts (see Table 25-4). The regressive hypothesis is also supported by the Mimivirus. The investigators who sequenced the Mimivirus genome reported that its genome is larger than the genomes of 20 different bacteria and archaea. Also unlike other viruses, Mimivirus has genes that encode components for protein translation. These researchers suggest that this giant virus evolved from a more complex cellular ancestor and gradually lost some of the genes necessary for protein synthesis. This hypothesis explains how viruses could have existed before their present-day hosts evolved. Recent studies on the origin of polydnaviruses contribute to our understanding of viral evolution. Polydnaviruses are particles that consist of multiple circles of double-stranded DNA encased in capsid proteins and an envelope. Each circle of DNA contains part of the virus genome. These viruses are found in ovary cells of many species of parasitic wasps. The polydnaviruses are unusual in that their circular DNA does not have genes for making the proteins needed to replicate and produce new viruses. Genes needed for viral replication are found in the wasp genome; the viruses can replicate only in the wasp ovary cells. The wasp injects polydnaviruses along with her eggs into certain caterpillars. The polydnaviruses express toxins that interfere with the caterpillar’s immune defenses and development. The wasp eggs hatch and develop inside the caterpillar, feeding on the caterpillar. Biologists wondered whether the polydnaviruses were really viruses. Were the polydnaviruses simply particles derived from wasp genes? An alternative hypothesis was that millions of years ago a particular virus infected wasps. The virus genome became integrated into the wasp genome and lost the capacity to enter virus particles. Instead, wasp DNA was incorporated into the particles. Read the Key Experiment, FIGURE 24-8, to learn which hypothesis is supported by the recent research of Jean-Michel Drezen, Annie Bézier, and their colleagues. No matter how viruses came to be and how they continue to evolve, they are here in large numbers and are important players in evolution. They mutate constantly, producing many new alleles. They replicate rapidly, and some of their genes are incorporated into the genomes of host cells, an example of horizontal gene transfer. In fact, viruses may be a major source of new genes for their host cells.

Review ■

■

Which of the hypotheses presented do you think best explains the origin of viruses? Why? In what way are polydnaviruses mutualistic partners with certain wasps?

24.6 SUBVIRAL AGENTS

KEY EXPERIMENT

■ ■ LEARNING OBJECTIVE

QUESTION: What is the origin of polydnaviruses?

9 Compare and contrast satellites, viroids, and prions.

infected wasps.

Subviral agents are infective agents that are smaller and simpler than viruses. Subviral agents include satellites, viroids, and prions.

EXPERIMENT: Annie Bézier, Jean-Michel Drezen, and their colleagues searched for virus-related genes in the wasp genome. The team analyzed DNA from three different wasp species. They compared nucleotide sequences with sequences of public databases containing sequences of all organisms and viruses studied.

Satellites depend on helper viruses

HYPOTHESIS: Polydnaviruses evolved from ancient viruses that

Satellites are subviral agents that depend on co-infection of a host cell with a helper virus. The satellite, which consists of either DNA or RNA, cannot reproduce without the helper virus. After a satellite encodes the protein for its protein coat, it is called a satellite virus. The agent that causes hepatitis D is a satellite that can reproduce only when the hepatitis B virus is also present. Sputnik is a satellite that infects the Mimivirus, which in turn infects an amoeba. Sputnik is dependent on the replication and assembly machinery set up by Mimivirus. Some researchers now refer to Sputnik as a virophage.

Viroids are the smallest known pathogens In 1971, Theodor Otto Diener, a plant pathologist, discovered an infective agent in potatoes that he named a viroid. Since that time, more than 30 species have been identified. Smaller than a virus, a viroid consists of a very short, circular, single strand of naked RNA (only 250 to 400 nucleotides). The viroid has no protective protein coat and no associated proteins to assist in duplication. Its RNA serves as a template that is copied by host RNA polymerases. Viroids are extremely hardy and resist heat and ultraviolet radiation because of the condensed folding of their RNA. Many viroids have been identified that cause plant diseases, including potato spindle tuber viroid, apple scar skin viroid, avocado sunblotch viroid, and tomato chlorotic dwarf viroid. Viroids cause stunted or distorted growth and sometimes kill the plant. For example, cadang-cadang disease has killed more than 30 million coconut palms trees in the Philippines. Viroids are transmitted by infected pollen or seeds. Viroids are generally found within the host-cell nucleus and appear to interfere with gene regulation. Molecular biologists have hypothesized that viroids mirror gene sequences in their hosts and silence critically important host genes. The viroid’s single-stranded RNA replicates, forming double-stranded RNA. The plant’s defense response cleaves this viroid RNA, producing small interfering RNAs (siRNAs); discussed in Chapter 13. However, the viroid siRNAs then cause host ribonucleases to selectively cleave host mRNAs with complementary base sequences. This action inactivates host mRNA and silences specific host genes. The viroids, themselves, are resistant to RNA silencing.

Prions are protein particles Stanley Prusiner, professor of neurology and biochemistry at the University of California School of Medicine, San Francisco, began his studies of prions in the early 1970s, motivated by the death

Juline Herbinière

Polydnavirus particles

250 nm

RESULTS AND CONCLUSION: The researchers identified 22 genes in wasp ovaries where viral particles are produced. These genes are present in the wasp genome and resemble key genes of nudiviruses, a family of insect viruses. The investigators found that many of these genes code for proteins that are part of the polydnavirus particles. These genes were conserved in several different wasp families that have polydnaviruses. The research team concluded that the polydnavirus evolved from a nudivirus that infected wasps millions of years ago. In time, the virus genome became incorporated into the wasp genome. Proteins needed for viral replication are now part of the wasp’s DNA, and the virus can replicate only in the wasp’s ovaries. At the same time, the wasps depend on the virus because the virus depresses the caterpillar’s immune defenses, allowing the wasp larvae to develop in the caterpillar. Thus, a mutualistic relationship has evolved between the wasp and the polydnavirus. The investigators suggested that the polydnavirus acts as a gene vector, transporting large chunks of DNA to the caterpillar. Might learning more about this mechanism have clinical applications? Source: Bézier, Annie, et al., “Polydnaviruses of Braconid Wasps Derive from an Ancestral Nudivirus.” Science, Vol. 323, No. 5916, Feb. 13, 2009, pp. 926–930.

FIGURE 24-8 Exploring the evolution of polydnaviruses The TEM shows polydnavirus particles in the nucleus of a cell in a wasp ovary. The polydnavirus particles are located at the periphery of an electron-dense region thought to be a “viral factory.”

of a patient from Creutzfeldt–Jakob disease (CJD), a degenerative brain disease. Prusiner discovered that the infective agent was not affected by radiation (which typically mutates nucleic acids), and he could not find DNA or RNA in the particles. In 1982, he named the infective agent prion, for “proteinaceous infectious particle.” Pruisner’s hypothesis that an infectious agent could cause disease without nucleic acids was not readily accepted because there was no known mechanism by which this could occur. However,

Pruisner and other researchers continued to study prions and discovered how they can cause cells to malfunction. In 1997, Prusiner was awarded the Nobel Prize in Physiology or Medicine for his discovery of prions—a new biological principle of infection. Prusiner and others have shown that animals have a gene that encodes a normally harmless protein known as PrP. Normal forms of the protein are found on the surfaces of brain cells and many other types of cells. The PrP protein, which consists of 208 amino acids, helps neurons transport copper and may play a role in the sense of smell in animals. Recently, researchers have discovered that normal prion proteins interact with the peptides associated with Alzheimer’s disease. Sometimes, the PrP protein folds into a different shape, an insoluble form that can cause disease. This misfolded protein is the prion. Mutations in the gene that encodes the PrP protein increase the risk that the protein will misfold and become a prion. The prion then somehow induces other PrP molecules to misfold into the pathogenic form (FIG. 24-9). Prions apparently can aggregate and accumulate in the brain and in certain other tissues and cause serious damage. Prions are found in the brains of patients with transmissible spongiform encephalopathies (TSEs). This group of fatal degenerative brain diseases has been identified in birds and mammals. These diseases are called TSEs because when infected, the brain appears to develop holes and becomes somewhat spongelike. The TSEs are in some ways like viral illnesses. However, there is a long latency period (about 5 years in cows and up to 10 years in humans) between contracting the infection and developing symptoms of the illness. Also, the TSEs have proven very difficult to treat. Genetically engineered mice that lack the prion protein gene are immune to TSE infection. The oldest known prion disease is scrapie in sheep and goats. When infected, animals lose coordination, become irritable, and itch so severely that they scrape off their wool or hair. Bovine spongiform encephalopathy (BSE) is a related prion disease popularly referred to as “mad cow disease” because some diseased cattle become aggressive. BSE became epidemic in cattle in the United Kingdom in the 1990s. More than 150 people have died from a human variety of BSE, providing evidence that the disease is transmissible from cow to human. The human disease is called vCJD because it is a variant of CJD, which is caused by the transformation of PrP proteins into prions. The infectious agent of vCJD has been recovered from infected human neural tissue and appears similar to the prion that causes scrapie in sheep. New regulations have been instituted in several countries to safeguard the food supply. Human-to-human transmission of vCJD has been associated with tissue and organ transplants and transfusion with contaminated blood. Chronic wasting disease, an illness related to mad cow disease, has spread among deer and elk populations in North America.

Contacts Prion

Contacts

Normal protein (PrP) 1

Prion induces normal PrP to misfold, forming another prion.

2

Each prion can induce additional PrP proteins to misfold.

3

Proteins aggregate.

Contacts

FIGURE 24-9 A model for how prion populations expand Prions contact normal PrP proteins and induce them to misfold and become prions. Each new prion can then contact additional PrP molecules and induce them to misfold, thus expanding the prion population. Prions can form clumps by aggregating.

Studies are under way to determine whether this disease can infect livestock or humans. A disease that is killing up to 70% of cheetahs in captivity also appears to be a prion disease. In this disease, large deposits of a misfolded protein accumulate in the liver and spleen. Like viruses, prions exhibit variability. There are different strains of prions that may differ from one another in the conformation of their proteins, in their properties, and in the diseases they cause. Prions are different from viruses in that they can arise spontaneously, most likely as a result of mutation. Researchers are studying ways to block prions from forming. They are also looking for ways to stimulate cells to destroy prions. Early diagnosis is critical, and the search is on for a blood test that would detect prion disease.

Review ■ ■

How do viroids differ from prions? How do viroids and prions differ from viruses?

■ ■

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

24.1 (page 502)

■

1 Contrast a virus with a cellular organism. A virus is a very small infective agent that consists of a core of nucleic acid and is dependent on a living host. Viruses contain the nucleic acids necessary to make copies of themselves, but in order to reproduce they must invade living cells and commandeer their metabolic machinery. 2 Describe the structure of a virus. ■ A virus is a subcellular particle consisting of a DNA or RNA genome surrounded by a protein coat, which is called a “capsid.” Some viruses are surrounded by an outer envelope. ■

Learn more about virus structure by clicking on the figure in CengageNOW.

24.2 (page 504) 3 Identify three characteristics used to classify viruses. ■

Viruses are classified based on host range, the types of organisms they infect; the type of nucleic acid they contain; and whether the nucleic acid is single-stranded or double-stranded. Other factors used to classify viruses include size and shape, presence of an envelope, and method of transmission from host to host.

24.3 (page 504) 4 Characterize bacteriophages. ■

Bacteriophages, or phages, are complex viruses that infect bacteria. They typically consist of a long molecule of dsDNA coiled within a polyhedral head.

The nucleic acid of some phages becomes integrated into the bacterial DNA and is then called a prophage. Bacterial cells that carry prophages are lysogenic cells. In lysogenic conversion, bacterial cells containing certain temperate viruses exhibit new properties. Watch the lytic and lysogenic cycles by clicking on the figure in CengageNOW.

24.4 (page 506) 6 Contrast viral infection of plants and animals, and identify specific diseases caused by viruses. ■ Most plant viruses are ssRNA viruses that do not have envelopes. Plant viruses can be spread among plants by insect vectors. Viruses spread through the plant via plasmodesmata. ■ Viruses enter animal cells by membrane fusion or by endocytosis. Viral nucleic acid is replicated within the host cell, proteins are synthesized, and new viruses are assembled and released from the cell. Among the diseases caused by DNA viruses are smallpox, herpes, respiratory infections, and gastrointestinal disorders. RNA viruses cause influenza, upper respiratory infections, AIDS, and some types of cancer. 7 Describe the reproductive cycle of a retrovirus, such as human immunodeficiency virus (HIV). ■ Retroviruses, such as HIV, use reverse transcriptase to transcribe their RNA genome into a DNA intermediate that becomes integrated into the host DNA. Copies of the viral RNA are then synthesized. Watch the HIV life cycle by clicking on the figure in CengageNOW.

Phage

24.5 (page 512) Phage DNA

Bacterium Bacterial DNA

Lytic cycle

Lysogenic cycle

8 Trace the evolutionary origin of viruses according to current hypotheses and describe recent research regarding the evolution of polydnaviruses. ■ According to the cellular origin hypothesis, viruses may be bits of nucleic acid that originally “escaped” from cellular organisms. ■ According to the coevolution hypothesis, viruses evolved before the three domains diverged; biologists who support this hypothesis think it unlikely that the similarities between viruses that infect archaea and those that infect bacteria evolved twice. ■ According to the regressive hypothesis, viruses evolved from independent cells that were parasites of larger cells. ■ Recent research suggests that polydnaviruses evolved from viruses that infected wasps. They have become mutualistic partners with their wasp hosts.

5 Contrast a lytic cycle with a lysogenic cycle. ■

■

In a lytic cycle, the virus destroys the host cell. There are five steps in a lytic cycle: attachment to the host cell; penetration of viral nucleic acid into the host cell; replication of the viral nucleic acid; assembly of newly synthesized components into new viruses; and release from the host cell. The genome of a temperate virus can be replicated along with replication of the host’s DNA without causing death of the host cell. This is a lysogenic cycle.

24.6 (page 513) 9 Compare and contrast satellites, viroids, and prions. ■

Satellites, viroids, and prions are smaller than viruses. Satellites reproduce only with the help of a helper virus. A viroid consists of a short strand of RNA with no protein coat. Many viroids cause plant diseases. A prion consists only of protein. Prions cause transmissible spongiform encephalopathies (TSEs).

T E S T YO U R U N D E R S TA N D I N G 1. The genome of a virus consists of (a) DNA (b) RNA (c) prions (d) DNA and RNA (e) DNA or RNA 2. The capsid of a virus consists of (a) protein subunits (b) nucleic acid (c) helical lipids (d) a carbohydrate envelope (e) RNA and lipid 3. Viruses that kill host cells are (a) lysogenic (b) lytic (c) viroids (d) prophages (e) temperate 4. Arrange the following list into the correct sequence for viral reproduction. 1. penetration 2. assembly 3. replication 4. attachment 5. release (a) 1, 2, 3, 4, 5 (b) 5, 2, 3, 4, 1 (c) 4, 1, 3, 2, 5 (d) 4, 1, 2, 3, 5 (e) 3, 1, 2, 4, 5 5. In lysogenic conversion (a) bacterial cells may exhibit new properties (b) the host cell dies (c) prions sometimes convert to viroids (d) reverse transcriptase transcribes DNA into RNA (e) lytic viruses become temperate 6. The types of organisms a particular virus can infect are referred to as its (a) assembly (b) commensal cohort (c) host range (d) prophage range (e) restriction cycle 7. Plant viruses (a) typically cause rapid death of infected plants (b) are mainly retroviruses (c) are killed with antibiotics (d) are transmitted throughout the plant body by insects (e) can be transmitted from one generation to the next through infected seeds

somal vesicle forms (d) viral DNA replicates in the host cytoplasm (e) the viral envelope fuses with the host nuclear membrane 9. Arrange the following list into the correct sequence for part of the cycle of a retrovirus. 1. dsDNA integrated into host DNA 2. viral proteins synthesized on host ribosomes 3. viral DNA uses host enzymes to transcribe viral RNA 4. reverse transcriptase catalyzes synthesis of ssDNA 5. synthesis of complementary DNA strand (a) 5, 2, 1, 3, 4 (b) 5, 2, 3, 4, 1 (c) 4, 5, 1, 3, 2 (d) 4, 1, 2, 3, 5 (e) 2, 1, 3, 4, 5 10. According to the cellular origin hypothesis, viruses (a) appeared before the three domains diverged (b) came from mobile genetic elements and had multiple origins (c) evolved from early plant cells (d) are bits of nucleic acid that escaped from cellular organisms (e) evolved from cells that were parasites in larger cells 11. Prions (a) consist of RNA with no protein coat (b) are misfolded proteins (c) cause several important plant diseases (d) are the cause of several important emerging diseases (e) consist of proteins that stimulate host RNA to produce DNA 12. What does this diagram illustrate? Complete the labels.

8. When a virus enters an animal cell by membrane fusion (a) the viral envelope fuses with the host plasma membrane (b) the capsid remains intact until the virus is released from the cell (c) an endo-

CRITICAL THINKING 1. How are viral genomes different from those of plant or animal cells? 2. How do viroids and prions differ? In what ways are both different from viruses? 3. EVOLUTION LINK. Discuss the coevolution of organisms that are mutualistic partners. Hint: You may want to use the example of polydnaviruses. (See discussion and Key Experiment on polydnaviruses.) 4. EVOLUTION LINK. Historically, biologists thought that viruses, because of their simple structure, evolved before cellular organisms. Later, the hypothesis that viruses escaped from cells gained

favor. A third view is the regressive hypothesis. Based on what you have learned about viruses, present an argument for one of these hypotheses. 5. SCIENCE, TECHNOLOGY, AND SOCIETY. What is the continuous challenge that influenza virus presents to scientists and public health officials? How can they meet this challenge? Additional questions are available in CengageNOW at www.cengage.com/ login.

25

Bacteria and Archaea

K. Lounatmaa/Photo Researchers, Inc.

Color-enhanced TEM of a bacterium (Streptococcus pyogenes) dividing by binary fission. This bacterium, a pathogen that inhabits the human nose and throat, can cause scarlet fever and inflammation of the heart tissue. The strain shown here is resistant to antibiotics, and infection can be fatal.

KEY CONCEPTS

I

n this chapter, we examine the diversity and characteristics of bacteria and archaea, two of the three main branches of the tree of life. These

25.1 Bacteria and archaea are single-celled organisms that, in contrast to eukaryotic cells, do not have membrane-enclosed organelles; most have a cell wall that surrounds the plasma membrane, and have a single, circular DNA molecule.

organisms, informally called prokaryotes, have inhabited our planet for

25.2 Evolution occurs rapidly in prokaryotes; natural

mately half of Earth’s biomass, the mass of living material. In contrast,

more than 3.5 billion years—much longer than eukaryotes, which evolved about 2.2 billion years ago. Although prokaryotes are microscopic, they are so numerous that they, together with the fungi, account for approxi-

selection acts on the genetic variation provided by mutations and genetic recombination, and is facilitated by rapid reproduction.

plants account for about 35% and animals for about 15% of the biomass of

25.3 The great diversity that has evolved in their metabo-

and other microorganisms in 1674 when he looked at a drop of lake water

lism and mode of nutrition allows prokaryotes to thrive in all kinds of habitats.

25.4 Prokaryotes make up two of the three domains:

our planet. The Dutch microscopist Anton van Leeuwenhoek discovered bacteria through a glass lens. During the late 1800s, many microorganisms, including some bacteria, fungi, and protozoa, were identified as pathogens,

Bacteria and Archaea.

agents that cause disease (see photograph). Bacteria cause many dis-

25.5 Prokaryotes play critical roles in ecology, commerce, and technology.

eases, for example, tuberculosis, tetanus, respiratory infections, and food

25.6 The many adaptations that have evolved in patho-

are pathogens. In fact, bacteria play essential roles in the biosphere. As

genic bacteria contribute to their great success.

decomposers, they break down organic molecules into their components.

poisoning in humans. However, only a small minority of bacterial species

Along with fungi, prokaryotes are nature’s chief recyclers. Without these microorganisms, elements such as carbon, nitrogen, phosphorus, and sulfur would remain locked up in the wastes and dead bodies of plants, animals, and other organisms and would be unavailable for the synthesis of new cells and organisms. Some prokaryotes are producers that carry on photosynthesis. Others convert atmospheric nitrogen to ammonia and then to nitrates, forms that are used by plants (see Fig. 55-8). This conver-

otes is fundamentally different from that of the eukaryotic cells of other living organisms.) Microbiologists assign prokaryotes to two domains: Archaea and Bacteria. Most prokaryotic cells are very small. Typically, their diameter ranges from 0.5 to 1.0 mm, and their length, from 1.0 to 5.0 mm. Their cell volume is only about one-thousandth that of small eukaryotic cells, and their length is only about one-tenth. Most prokaryotes are unicellular, but some form colonies or filaments containing specialized cells.

sion enables plants and animals (because they eat plants) to manufacture essential nitrogen-containing compounds such as proteins

Prokaryotes have several common shapes

and nucleic acids.

Two basic prokaryote shapes are spherical and rod-shaped. Spherical prokaryotes, known as cocci (sing., coccus), occur singly in some species and in groups of independent cells in others (FIG. 25-1a). Cells may be grouped in twos (diplococci), in long chains (streptococci), or in irregular clumps that look like bunches of grapes (staphylococci). Rod-shaped prokaryotes, called bacilli (sing., bacillus), may occur as single rods or as long chains of rods (FIG. 25-1b). Some prokaryotes form spirals. If the spiral-shaped cell is flexible, it is a spirochete; if rigid, it is a spirillum (pl., spirilla) (FIG. 25-1c). A spirillum shaped like a comma is called a vibrio. Some archaea have unusual shapes, such as triangular or square-shaped cells.

We begin this chapter with a description of the structure of bacteria and archaea, and then discuss prokaryote reproduction and evolution. We outline some of the nutritional and metabolic adaptations of these organisms, and then describe the phylogeny of the two prokaryote domains. We present an overview of the impact of prokaryotes on ecology, technology, and commerce, and end the chapter with a discussion of bacteria and disease.

25.1 THE STRUCTURE OF BACTERIA AND ARCHAEA ■ ■ LEARNING OBJECTIVES 1 2 3

Describe the structure and common shapes of prokaryotic cells. Contrast the bacterial cell wall in gram-positive and gram-negative bacteria. Describe movement in prokaryotes, and describe the structure of the bacterial flagellum.

Center for Disease Control

0.6 μm

In contrast to eukaryotic cells, prokaryotic cells do not have a nucleus or other membrane-enclosed organelles (FIG. 25-2; see Chapter 4). Although the prokaryotic cell does not have a membrane-enclosed nucleus, it does have a nuclear area, also referred to as the nucleoid, which contains DNA. The dense cytoplasm of the prokaryotic cell contains ribosomes (smaller than those found in eukaryotic cells) and storage granules that hold glycogen, lipid, and phosphate compounds. Enzymes needed for metabolic activities may be located in the cytoplasm. Although the membranous organelles of eukaryotic

© Dr. Gary Gaugler /Photo Researchers, Inc.

© Dennis Kunkel Microscopy / Visuals Unlimited, Inc.

In contrast to viruses, viroids, and prions, which consist only of nucleic acid and/or protein, prokaryotes are cellular organisms. (Recall from Chapters 4 and 23 that the cell structure of prokary-

Prokaryotic cells do not have membrane-enclosed organelles

3 μm

(a) Cocci bacteria. Colorized SEM of Staphylococcus aureus. These bacteria cause skin and wound infections, food poisoning, and toxic shock syndrome.

FIGURE 25-1 Common shapes of prokaryotes

(b) Bacilli bacteria. Colorized SEM of Salmonella. These bacilli cause food poisoning.

12 μm

(c) Spirochete bacteria. Colorized SEM of Borrelia burgdorferi. These spirochetes cause Lyme disease, transmitted by infected deer ticks.

KEY POINT

In contrast to eukaryotic cells, prokaryotic cells do not have a nucleus or other organelles enclosed by membranes. They typically have a nuclear area with a single, circular DNA molecule.

Outer membrane Fimbriae (structures used for attachment)

Peptidoglycan layer

Cell wall

Nuclear area (nucleoid)

Storage granule Plasmid (DNA)

Flagellum Ribosomes Bacterial chromosome (DNA)

Capsule Plasma membrane

FIGURE 25-2 Animated Structure of a prokaryotic cell This bacillus is a gram-negative bacterium (discussed in text). Note the absence of a nuclear envelope surrounding the bacterial DNA.

cells are absent, in some prokaryotic cells the plasma membrane is extensively folded inward. Enzymes needed for cellular respiration and photosynthesis may be associated with the plasma membrane or its folds.

A cell wall protects most prokaryotes Most prokaryotic cells have a cell wall surrounding the plasma membrane. The cell wall provides a rigid framework that supports the cell and maintains its shape. Bacterial cells have a high concentration of dissolved solutes. The cell wall keeps the cell from bursting under hypotonic conditions (see Chapter 5). Thus, most bacteria are adapted to hypotonic surroundings. When wall-less forms of bacteria are produced experimentally, they must be maintained in isotonic solutions to keep them from bursting. However, cell walls are of little help when a bacterium is in a hypertonic environment, as in food preserved by a high sugar or salt content. For this reason, most bacteria grow poorly in jellies, jams, salted fish, and other foods preserved in these ways. The bacterial cell wall includes peptidoglycan, a complex polymer that consists of two unusual types of sugars (amino sugars) linked with short polypeptides. The sugars and polypeptides

are joined to form a single macromolecule that surrounds the entire plasma membrane. Peptidoglycan is absent in the archaean cell wall. Differences in bacterial cell wall composition are of great interest to microbiologists and are important clinically. In 1888, the Danish physician Christian Gram developed the Gram staining procedure. Bacteria that absorb and retain crystal violet stain in the laboratory are referred to as gram-positive, whereas those that do not retain the stain when rinsed with alcohol are gram-negative. The cell walls of gram-positive bacteria are very thick and consist primarily of peptidoglycan. The cell walls of gram-negative bacteria have two layers: a thin peptidoglycan layer and a thick outer membrane. The outer membrane resembles the plasma membrane but contains polysaccharides bonded to lipids (FIG. 25-3). Distinguishing between gram-positive and gram-negative bacteria is important in treating certain diseases. For example, the antibiotic penicillin interferes with peptidoglycan synthesis, ultimately resulting in a fragile cell wall that cannot protect the cell (see Chapter 7 discussion of drugs that are enzyme inhibitors). Predictably, penicillin works most effectively against grampositive bacteria.

Some bacteria produce capsules or slime layers Many prokaryote species produce a capsule or slime layer that surrounds the cell wall. A slime layer is more loosely attached to the cell wall than a capsule. These outer layers are made of polysaccharide or protein. In free-living species, the outer covering may provide the cell with added protection against phagocytosis (engulfment; see Chapter 5) by other microorganisms. In disease-causing bacteria, a capsule or slime layer may protect against phagocytosis by the host’s white blood cells. The ability of Streptococcus pneumoniae to cause bacterial pneumonia depends on its capsule. A strain of S. pneumoniae that lacks a capsule does not cause the disease. Bacteria also use their capsules to attach to surfaces such as rocks, plant roots, or human teeth (where they cause dental plaque).

Some prokaryotes have fimbriae or pili Some prokaryotes have hundreds of hairlike appendages called fimbriae (sing., fimbria). Fimbriae, which are made of protein, are shorter than flagella. Pili (sing., pilus) are usually longer than fimbriae. There are typically fewer pili on the cell surface than fimbriae. Bacteria use fimbriae and pili to attach to cell surfaces,

Alfred Pasieka/Peter Arnold, Inc.

Thick peptidoglycan layer

Cell wall

Plasma membrane Endospores

Transport protein

(a) Gram-positive cell wall. A thick layer of peptidoglycan molecules is held together by amino acids.

Polysaccharides Lipoprotein

Outer membrane

Cell wall

Thin peptidoglycan layer Plasma membrane Transport protein

(b) Gram-negative cell wall. A thin layer of peptidoglycan is covered by an outer membrane.

FIGURE 25-3 Bacterial cell walls

including the surfaces of cells they infect. These protein structures also help bacteria adhere to one another. Some elongated pili, called sex pili, are important in transmitting DNA between bacteria.

Some bacteria survive unfavorable conditions by forming endospores When the environment becomes unfavorable, for example, when nutrients are limited or the environment becomes very dry or

2 μm

FIGURE 25-4 Endospores Color-enhanced TEM of Clostridium tetani, the bacterium that causes tetanus. Each bacterial cell (blue) contains one endospore (orange), a resistant, dehydrated cell that develops within the original cell.

hot, some types of bacteria become dormant. The cell loses water, shrinks slightly, and remains inactive until water is again available. Certain types of bacteria form dormant, extremely durable cells called endospores. After an endospore forms, the cell wall of the original cell lyses, releasing the endospore. Formation of endospores is not a type of reproduction in bacteria; endospores are not comparable to the reproductive spores of fungi and plants. Only one endospore is formed per original cell, so the total number of individuals does not increase. Archaea do not form endospores; these prokaryotes produce unique enzymes on the cell surface that protect them from cold, heat, and desiccation. Endospores survive in very dry, hot, or frozen environments or at times when food is scarce (FIG. 25-4). Some endospores are so resistant that they can survive an hour or more of boiling or centuries of freezing. When environmental conditions are again suitable for growth, the endospore germinates, forming an active, growing bacterial cell. Several types of bacteria that form endospores cause disease. The endospore of Bacillus anthracis, the bacterium that causes anthrax, is so hardy that this pathogen has become a concern as an agent of biological warfare. The bacteria (Clostridium tetani) that cause tetanus and the bacteria (C. perfringens) that cause gas gangrene typically enter the body with soil when a person suffers a deep cut or puncture wound. Patients may also be exposed to these serious diseases when surgical instruments are not effectively sterilized, allowing endospores to survive.

Many types of prokaryotes are motile Can you imagine trying to swim through molasses? Water has the same relative viscosity to prokaryotes that molasses has to humans. Most motile prokaryotes manage to move by means of rotating

Eye of Science/ Photo Researchers, Inc.

flagella. The number and location of flagella are important in classifying some bacterial species. Unlike eukaryotic flagella, prokaryotic flagella do not consist of microtubules (see Chapter 4). A bacterial flagellum is a long, thin appendage, consisting of three parts: a basal body, a hook, and a single filament (FIG. 25-5). The basal body is a complex structure that anchors the flagellum into the cell wall by disc-shaped plates. The curved hook connects the basal body to the long, hollow filament that extends into the outside environment. The basal body is a motor, somewhat like the propeller of a boat motor. The bacterium uses energy from ATP to pump protons out of the cell. Diffusion of these protons back into the cell powers the motor that spins the flagellum. The rotary motion produced pushes the cell much like a propeller pushes a ship through the water. The flagellum rotates counterclockwise, pushing the cell forward. Archaea also have flagella. These flagella are thinner than those of bacteria. Biologists do not yet know the structure of the motor in the archaeal flagellum. Some prokaryotes that lack flagella move by gliding. Many prokaryotes exhibit chemotaxis, movement in response to chemicals in the environment. For example, bacteria move toward food. Certain bacteria also move toward one another. However, bacteria move away from certain harmful chemicals.

0.5 μm

(a) Color-enhanced TEM of Vibrio cholerae. This flagellate bacterium causes cholera in humans.

Review ■ ■

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In what ways do prokaryotic cells differ from eukaryotic cells? How do gram-positive and gram-negative bacteria differ? Why is this important? How does a bacterial flagellum differ from a eukaryotic flagellum?

Plasma membrane

25.2 PROKARYOTE REPRODUCTION AND EVOLUTION

Cytoplasm

■ ■ LEARNING OBJECTIVES Basal body

mechanisms (transformation, transduction, and conjugation) that may lead to genetic recombination. State specific factors that contribute to the rapid evolution of bacteria.

The genetic material of a prokaryote lies in the nuclear area, but is not surrounded by a nuclear envelope. In most species, the genetic material is contained in a single, circular DNA molecule. If stretched out to its full length, this molecule would be about 1000 times as long as the cell itself. Unlike the DNA in eukaryotic chromosomes, prokaryote DNA has little protein associated with it. In addition to their genomic DNA, most bacteria have one or more plasmids, smaller circular fragments of DNA. Bacterial plasmids often have genes that code for catabolic enzymes, for genetic exchange, or for resistance to antibiotics. Plasmids replicate independently of the genomic DNA or become integrated into it (see Chapter 15).

Outer membrane

Protein rings

4 Describe asexual reproduction in prokaryotes, and summarize three

5

Peptidoglycan layer

Hook

Cell wall

Filament

(b) Structure of a bacterial flagellum. The basal body is the motor. It consists of a series of disc-shaped plates that anchor the flagellum to the cell wall and plasma membrane. These plates spin the hook and filament of the flagellum.

FIGURE 25-5 Bacterial flagella

Rapid reproduction contributes to prokaryote success Prokaryotes are extremely successful organisms in terms of their numbers and distribution. Their success is in large part due to their remarkable ability to reproduce rapidly. Prokaryotes reproduce asexually, generally by binary fission, a process in which one cell divides into two similar cells, as in the chapter opening photograph (also see Fig. 10-12). First, the circular DNA replicates; and then an ingrowth of both the plasma membrane and the cell wall forms a transverse wall. Binary fission occurs with remarkable speed. Under ideal conditions, some bacterial species divide in less than 20 minutes. At this rate, if nothing interfered, one bacterium could give rise to more than one billion bacteria within 10 hours! Fortunately, bacteria cannot reproduce at this rate for very long before lack of food or the accumulation of waste products slows their population expansion. A less common form of asexual reproduction among bacteria is budding. In budding, a cell develops a bulge, or bud, that enlarges, matures, and eventually separates from the mother cell. A few species of bacteria (actinomycetes) divide by fragmentation. Walls develop within the cell, which then separates into several new cells.

Prokaryotes transfer genetic information Although sexual reproduction involving the fusion of gametes does not occur in prokaryotes, genetic material can be exchanged among bacteria and among archaea. Such exchange of genes, or gene transfer, results in genetic recombination. Horizontal gene transfer occurs when an organism transfers genetic material to another organism that is not its offspring. (Transfer of genetic mate-

1

2

3

DNA exchanged

rial from parent to offspring is called vertical gene transfer.) Gene transfer among bacteria takes place by three different mechanisms: transformation, transduction, and conjugation. 1. When bacteria die, they release DNA that can be taken in by certain other bacteria. In transformation, a bacterial cell takes up fragments of foreign DNA (or RNA) released by another bacterium. The DNA must bind to DNA-binding proteins on the surface of the bacterium. Once it enters the bacterial cell, some of the DNA is incorporated into the host’s

Phage DNA with bacterial genes

1

DNA of a phage penetrates bacterial cell.

2

Phage DNA may become integrated with host-cell DNA as a prophage.

3

When the prophage becomes lytic, bacterial DNA is degraded and new phages are produced. New phages may contain some bacterial DNA.

4

Bacterial cell lyses and releases many phages, which can then infect other cells.

5

Phage infects new host cell.

6

Bacterial genes introduced into new host cell are integrated into host's DNA. They become a part of bacterial DNA and are replicated along with it.

Fragmented bacterial DNA

Bacterium dies and releases DNA.

Fragments of foreign DNA bind to proteins on surface of living bacterium.

DNA enters cell, and some DNA is incorporated into host cell by reciprocal recombination.

FIGURE 25-7 Transduction FIGURE 25-6 Transformation In transformation, foreign DNA from a bacterium that has died enters a host bacterium. Some DNA is exchanged, resulting in a recombinant cell.

In transduction, a phage transfers bacterial DNA from one bacterium to another, resulting in genetic recombination. Transduction is an important means of horizontal gene transfer.

Conjugation has been most extensively studied in the bacterium Escherichia coli. In the E. coli population, there are donor cells, or F+ cells, which have DNA that can be transmitted to recipient cells, or F− cells. F+ cells have a DNA sequence known as the F factor (F stands for fertility) that is necessary for a bacterium to serve as a donor during conjugation. The F factor can be in the form of a plasmid, or it can be part of the DNA in the bacterial chromosome. F genes encode enzymes essential for transferring DNA. Certain F genes encode sex pili, long, hairlike extensions that project from the cell surface. The sex pilus of an F+ cell recognizes and binds to the surface of an F− cell, forming a cytoplasmic conjugation bridge between the two cells. The F plasmid replicates itself, and DNA is transferred from donor to recipient bacterium through the conjugation bridge. F plasmids may also have other types of genes, including those that determine resistance to antibiotics.

Evolution proceeds rapidly in bacterial populations Because bacteria reproduce rapidly by binary fission, mutations are quickly passed on to new generations. Mutations that confer some advantage

KEY POINT

Conjugation, a mechanism of horizontal gene transfer, results in genetic recombination. Sex pilus

© Dennis Kunkel Microscopy, Inc./Phototake / Alamy

own genome by reciprocal recombination (the DNA is exchanged) between the new DNA and the host chromosome (FIG. 25-6). Recall from Chapter 12 that Oswald T. Avery and his colleagues identified DNA as the agent that transformed bacterial cells experimentally and showed that DNA is the chemical basis of heredity. Foreign DNA can also be taken up as plasmids. When this occurs, DNA does not undergo recombination; it remains as a plasmid separate from the bacterial chromosome. 2. In a different process of horizontal gene transfer, transduction, a phage carries bacterial genes from one bacterial cell into another (FIG. 25-7). Normally, a phage contains only its own DNA. However, sometimes a phage incorporates some of the bacterial DNA of its host. Then, when the phage infects another bacterium, it transfers that DNA to its new host. The bacterial chromosome of this new host then becomes a recombination of its own original DNA and the DNA from another bacterium. 3. In conjugation, two cells of different mating types come together, and genetic material is transferred from one to the other (FIG. 25-8). In contrast to transformation and transduction, conjugation involves contact between two cells.

0.5 μm

(a) Color-enhanced SEM of E. coli bacteria conjugating. The bacteria are connected by a sex pilus. When stimulated by the contact, the cells pull close together and form a conjugation bridge between donor and recipient cells (not shown). F+ (donor) cell

Bacterial chromosome

F– (recipient) cell 1

F+ (donor) cell produces sex pilus.

2

Sex pilus develops into conjugation bridge.

3

Single strand of F plasmid DNA is transferred from F+ cell to F– cell.

4

DNA replicates. Both bacterial cells now contain double-stranded F plasmid. The F– cell has been converted to an F+ cell.

F plasmid

(b) The process of conjugation.

FIGURE 25-8 Animated Conjugation In conjugation, a donor bacterium transfers plasmid DNA to a recipient bacterium. Conjugation requires cell-to-cell contact.

spread through the population, and the effects of natural selection are quickly evident. Horizontal gene transfer greatly contributes to the rapid evolution that takes place in prokaryotes. Acquisition of new DNA and genetic recombination are important sources of the genetic variation required for diversification and adaptation. New DNA introduced into a bacterium’s genome represents raw material for evolution. New genes are subject to mutation and are acted on by the forces of natural selection. The changes in the genetic material are passed to succeeding generations by binary fission. Changes that result in adaptation can quickly spread through future bacterial populations. Genetic recombination is also an important mechanism in the evolution of archaea.

Review ■

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How do prokaryotes reproduce? What mechanisms result in gene transfer? How does transduction contribute to the rapid evolution of bacterial populations? What are the steps that take place during conjugation?

25.3 NUTRITIONAL AND METABOLIC ADAPTATIONS ■ ■ LEARNING OBJECTIVE 6 Describe the principal modes by which prokaryotes carry on nutrition and energy capture, and compare their requirements for oxygen.

Based on the two main ways of obtaining nutrition, prokaryotes are classified as autotrophs (-troph comes from a Greek word that means “nutrition”) or heterotrophs. Autotrophs are able to use inorganic compounds, such as carbon dioxide, as a source of carbon for manufacturing their organic molecules. Most prokaryotes are heterotrophs that obtain carbon atoms from the organic compounds of other organisms. Based on two principal ways of capturing energy, autotrophs are classified as chemotrophs and autotrophs. Chemotrophs obtain their energy from chemical compounds. Phototrophs capture energy from light. More than three billion years ago, early bacteria de-

TABLE 25-1

veloped the ability to use the sun as an energy source. These early phototrophs used hydrogen sulfide to reduce carbon dioxide. They released sulfur as a waste product. About 2.7 billion years ago, the ability to use water, rather than hydrogen sulfide, to reduce carbon dioxide evolved in early cyanobacteria. Oxygen was released as a waste product. When we consider the source of carbon and the source of energy, we can classify prokaryotes into four main groups (TABLE 25-1): 1. Photoautotrophs, such as cyanobacteria, use the energy from sunlight to synthesize organic compounds from carbon dioxide and other inorganic compounds. 2. Chemoautotrophs also use carbon dioxide as a carbon source, but they do not use sunlight as their energy source. Instead, they obtain energy by oxidizing inorganic chemical substances such as ammonia (NH3) and hydrogen sulfide (H2S). 3. Photoheterotrophs, such as the purple nonsulfur bacteria, obtain their carbon from other organisms but use chlorophyll and other photosynthetic pigments to trap energy from sunlight. 4. The majority of prokaryotes are chemoheterotrophs. They depend on organic molecules for both their carbon and energy. Many bacterial chemoheterotrophs are free-living decomposers that obtain their carbon and energy from dead organic matter. These bacteria are sometimes called saprotrophs. Some chemoheterotrophs are pathogens, which obtain their nourishment from the organisms they infect. They harm their hosts by causing diseases. Other heterotrophic bacteria benefit their hosts. For example, some of the bacteria that inhabit the human large intestine produce vitamin K and certain B vitamins for their hosts.

Most prokaryotes require oxygen Whether they are heterotrophs or autotrophs, most bacterial cells are aerobic and require oxygen for cellular respiration. Many are facultative anaerobes that use oxygen for cellular respiration if it is available but can carry on metabolism anaerobically when necessary. Other bacteria are obligate anaerobes that carry out anaerobic respiration; they respire with terminal electron acceptors other than oxygen, for example, sulfate (SO42−), nitrate (NO3−), or iron (Fe2+). Some obligate anaerobes, including certain archaea, are actually killed by even low concentrations of oxygen.

Modes of Nutrition and Energy Capture

Mode of Nutrition Autotroph Photoautotroph Chemoautotroph

Heterotroph Photoheterotroph Chemoheterotroph

Energy Source

Carbon Source

Examples of Organisms

Sunlight Inorganic chemicals (e.g., NH3, H2S, Fe2+)

CO2 CO2

Cyanobacteria; purple sulfur bacteria Certain proteobacteria; most archaea, e.g., methanogens, extreme halophiles

Sunlight Organic compounds

Organic compounds Organic compounds

Purple and green nonsulfur bacteria Free-living decomposers; most bacterial pathogens

Domain Eukarya

Eukaryotes

Crenarchaeota

Spirochetes

Chlamydias

Domain Archaea Cyanobacteria

Delta

Gamma

Beta

Alpha

Epsilon

Proteobacteria

All organisms require nitrogen in order to manufacture amino acids and nucleic acids. Some bacteria (for example, certain cyanobacteria) and archaea (for example, methanogens) can reduce nitrogen in the atmosphere to ammonia. This process is called nitrogen fixation. Ammonia produced by nitrogen fixation is converted to ammonium ions (NH4+). Nitrogen-fixing bacteria can use these simple forms of nitrogen to produce organic compounds. Certain bacteria convert ammonia or ammonium ions to nitrite (NO2−), and others convert nitrite to nitrate (NO3−). This process, called nitrification, converts nitrogen to a form that can be used by plants and fungi. Animals obtain nitrogen from organic compounds when they eat other organisms. As we will discuss in a later section, all other organisms ultimately depend on nitrogen fixation and nitrification by prokaryotes for their survival (see Chapter 55).

Gram-positives

Domain Bacteria

Euryarchaeota

Some prokaryotes fix and metabolize nitrogen

Common ancestor of all living organisms

Review ■ ■

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How do chemoheterotrophs obtain energy? How do facultative anaerobes differ from obligate anaerobes? How do they differ from aerobes? How do bacteria obtain nitrogen needed to produce amino acids and nucleic acids?

25.4 THE PHYLOGENY OF THE TWO PROKARYOTE DOMAINS ■ ■ LEARNING OBJECTIVES 7 8

Compare characteristics of the three domains: Archaea, Bacteria, and Eukarya. Distinguish between the two main groups of archaea and identify specific types of archaea belonging to each group.

9 Describe the main groups of bacteria discussed in this chapter (see

FIGURE 25-9 Three domains This highly simplified diagram depicts some representative taxa of domain Bacteria and domain Archaea. The relationships illustrated are based on analyses of ribosomal RNA gene sequences. As taxonomists consider additional data, these relationships will be modified.

Based on Woese’s work and on other recent data, systematists now classify the modern descendants of these two ancient lines in two domains: Archaea and Bacteria (FIG. 25-9). These groups are thought to have diverged from a common ancestor about four billion years ago. Archaea and bacteria were the only living organisms on our planet for about two billion years. Thus, archaea and bacteria have had a very long time to evolve and adapt to all types of environments. Their diversity is staggering. Horizontal gene transfer has also contributed to the diversity of these organisms. As a result of gene transfer, sometimes from distantly related species, the genomes of archaea and bacteria are actually a mix of genes from many prokaryotes.

Table 25-4).

Under a microscope, most prokaryotes appear rather similar in size and form. However, using sequence analysis of small subunit 16S ribosomal RNA (SSU rRNA), Carl Woese and his co-workers demonstrated that there are two fundamentally different groups of prokaryotes (see Chapter 23). Each group has signature sequences, regions of SSU rRNA that have unique nucleotide sequences. The explanation is that after they diverged, prokaryote populations diversified and mutations occurred that affect RNA sequences. Using such analyses, Woese hypothesized that ancient prokaryotes split into two lineages early in the history of life.

Key characters distinguish the three domains Several key characters distinguish archaea from bacteria. In contrast to bacteria, archaea do not have peptidoglycan in their cell walls. Although their plasma membranes are structurally similar, they are chemically unique. In the plasma membranes of bacteria and eukaryotes, straight-chain fatty acids are linked to glycerol molecules by ester linkages. In contrast, fatty acid components are not found in archaea. Instead, branched-chain hydrocarbons (synthesized from isoprene units) are bonded to glycerol by ether linkages (TABLE 25-2).

TABLE 25-2

Comparison of the Three Domains

Characteristic

Bacteria

Archaea

Eukarya

Nuclear envelope Membrane-enclosed organelles Circular chromosome Number of chromosomes

Absent Absent Present (linear in some species) Typically one (may also have plasmid) Absent Present Straight-chain fatty acids bonded to glycerol by ester linkages 70S*

Absent Absent Present Typically one (may also have plasmid) Present Absent Branched-chain hydrocarbons linked to glycerol by ether linkages 70S

Present Present Absent Typically many

One relatively simple RNA polymerase Begins with formylmethionine Yes

One relatively complex RNA polymerase Begins with methionine Yes

Histones associated with DNA Peptidoglycan in cell wall Structure of lipids in plasma membrane Size of ribosomes RNA polymerase Translation Growth above 70°C

Present Absent Straight-chain fatty acids bonded to glycerol by ester linkages 80S except in mitochondria and chloroplasts Several relatively complex RNA polymerases Begins with methionine No

*The numbers 70S and 80S refer to the sedimentation coefficient (a measure of relative size) when centrifuged.

O R

C O R

Ester linkage

H R

C O R

H Ether linkage

The absence of a second electronegative oxygen atom makes ether linkages stronger than ester linkages. This unique membrane structure may contribute to the ability of the archaea to survive and thrive in harsh environments. In some ways, archaea are more like eukaryotes than like bacteria. For example, archaea do not have the simple RNA polymerase found in bacteria. Like eukaryotes, their translation process begins with methionine, whereas in bacteria, translation begins with formylmethionine (see Chapter 13). In addition, several antibiotics that affect bacteria do not affect archaea or eukaryotes. Bacteria also share some characters (that are absent in archaea) with eukaryotes. For example, as stated above, bacteria and eukarya both have ester-linked membrane lipids, whereas archaea have ether-linked membrane lipids. Some microbiologists hypothesize that eukaryotes are a product of fusion between an archaean, which contributed components for transcription and translation, and a bacterium, which contributed enzymes necessary for energy metabolism.

Taxonomy of archaea and bacteria continuously changes Prokaryote taxonomy is now based largely on molecular data, mainly RNA sequencing, and more recently, on sequencing entire genomes. Groups that branched off earlier had more time to accumulate mutations in their SSU rRNA. Their nucleotide sequences are less similar than those of groups that diverged more recently.

Microbiologists who are developing phylogenetic trees based on sequencing entire genomes argue that there may be 1000 genes that code proteins for every 1 gene that codes an rRNA. These researchers prefer to consider the proportions of genes (or proteins) that genomes of various groups have in common. Taxonomy of archaea and bacteria continuously changes as systematists study new molecular data that provide new clues to the phylogeny of these groups. Although prokaryotic taxonomy is controversial and continuously changing, about 7000 species of prokaryotes have been classified. Hundreds of thousands of additional species are thought to exist. The editors of Bergey’s Manual of Systematic Bacteriology, considered the definitive reference text by microbiologists, have divided archaea into 2 phyla and bacteria into about 20 phyla based on 16S rRNA analyses.

Many archaea inhabit harsh environments Domain Archaea consists of two major groups (phyla): Crenarchaeota and Euryarchaeota (TABLE 25-3). This phylogeny is based on 16S rRNA and on sequencing of entire genomes. Systematists have not yet determined whether these two groups represent phyla or higher level taxa. If they are higher level taxa, the branches of each group may represent phyla. The Crenarchaeota consist mainly of extreme thermophiles, archaea that require a very high temperature or very low temperature for growth. The optimum temperature for many is greater than 80°C (176°F). These archaea inhabit very hot, sometimes acidic, environments. One species is found in the hot sulfur springs of Yellowstone Park at temperatures near 80°C and pH values of 1 to 2, the pH of concentrated sulfuric acid (FIG. 25-10a). Others inhabit volcanic areas under the sea. One species, found near deepsea hydrothermal vents on the seafloor of the Pacific Ocean, lives at temperatures ranging from 80°C to 120°C. In contrast, some Crenarchaeota live in very cold environments (below 15°C).

© Helen E. Carr / Biological Photo Service

© Mike Cavaroc/ Alamy

(a) Extreme thermophiles. Orange and yellow colonies of extreme thermophiles thrive in the Grand Prismatic Spring in Yellowstone National Park in Wyoming, USA.

FIGURE 25-10 Archaea that inhabit extreme environments

Some of these archaea drift in the ocean and are an important part of the plankton. The Euryarchaeota also include many archaea that inhabit extreme environments. This group includes methanogens, extreme halophiles, and extreme thermophiles. The methanogens (methane producers) are a large, diverse group that inhabit oxygen-free environments in sewage, swamps, and the digestive tracts of humans and other animals. They are obligate anaerobes that produce methane gas from simple carbon compounds. The methanogens are important in recycling components of organic products of organisms that inhabit swamps. Methanogens that inhabit the digestive tracts of cows and other grazing animals produce methane, which is belched out by the animals. Methanogens produce more than 80% of the methane (more than two billion tons each year) in Earth’s atmosphere. Methane is an important greenhouse gas. Extreme halophiles are heterotrophs that require large amounts of Na+ for their growth. They live in saturated brine solutions such as salt ponds, the Dead Sea, and Great Salt Lake (FIG. 25-10b). The extreme halophiles use aerobic respiration to make ATP. However, they also carry out a form of photosynthesis in which they capture the energy of sunlight using a purple pigment (bacteriorhodopsin). This pigment is very similar to the pigment rhodopsin involved in animal vision. Nanoarchaeum is a very small (400 nm) anaerobic, extreme thermophile that is now classified as a euryarchaeote. This microbe

(b) Extreme halophiles. These seawater evaporating ponds near San Francisco Bay are colored pink, orange, and yellow from the large number of extreme halophiles that inhabit them. The colors are a result of pigments in the cell membranes. These bacteria are harmless, and the ponds are used to produce salt commercially.

is one of the smallest cellular organisms identified to date. It also has one of the smallest genomes (less than 500,000 nucleotides). Nanoarchaeum lives attached to another archaean and depends on its host for many of its metabolic needs. Many archaea inhabit less extreme conditions. For example, nonextreme archaea are abundant in the soil and in the cold ocean surface waters near Antarctica. Microbiologists have suggested that the archaea may be important in biogeochemical cycles and in marine food chains. No pathogenic archaea have been identified.

Bacteria are the most familiar prokaryotes The bacteria are widely distributed in the environment and are better known to microbiologists than are the archaea. Five major groups are considered here: proteobacteria (gram-negative), cyanobacteria (gram-negative), gram-positive bacteria, chlamydias (gram-negative), and spirochetes (gram-negative). These groups are summarized in TABLE 25-4.

Review ■

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What are two types of archaeans that inhabit extreme environments? What are methanogens? How do these bacteria differ (described in Table 25-4): (1) proteobacteria, (2) cyanobacteria, and (3) chlamydias?

TABLE 25-3

Two Groups of Archaea

Crenarchaeota Eye of Science / Photo Researchers, Inc.

1 μm

Some extremely thermophilic (thrive at 70°C, and some thrive at temperatures greater than 100°C); others can live at temperatures below 15°C. Many are chemoautotrophs. Important part of plankton in cold, oxygen-rich seas.

SEM of Pyrococcus furiosus, an anaerobe that inhabits marine sand This methanogen is highly resistant to heat; its optimum temperature is 100°C. Pyrococcus grows on sugars and small peptides, which serve as electron donors. DNA polymerase from P. furiosus is used in PCR (polymerase chain reaction) DNA amplification process because it is stable at high temperatures and is highly accurate. Euryarchaeota Dr. Kari Lounatmaa / Science Photo Library / Photo Researchers, Inc.

Includes methanogens, halophiles (not all extreme), and acidophiles. Some extreme thermophiles included in this phylum.

1 μm

TEM of a methanogen (Methanospirillum hungatii) undergoing cell division Two other archaea are visible in cross section. When not dividing, these archaea are spiral shaped.

TABLE 25-4

Major Groups of Bacteria

Science Photo Library

Proteobacteria (Gram-negative)

2 μm

SEM of Escherichia coli colony

© Phototake, Inc. / PhototakeUSA.com

E. coli are Gamma Proteobacteria.

50 μm

SEM of fruiting body of the myxobacterium Stigmatella aurantiaca Protective resting cells within the fruiting bodies are very resistant to heat and drying. Myxobacteria are Delta Proteobacteria.

A large, very diverse group of gram-negative bacteria. Based on rRNA sequences, the group is divided into five subgroups designated as alpha, beta, gamma, delta, and epsilon. Alpha Proteobacteria Includes many symbionts of plants and animals, and some pathogens. Rhizobium species live symbiotically in root nodules of legumes (e.g., beans) and convert atmospheric nitrogen to a form usable by plants (nitrogen fixation). Rickettsias are very small, rod-shaped bacteria. A few species are pathogenic to humans and other animals; transmitted by arthropods through bites or through contact with their excretions. Among these are typhus (transmitted by fleas and lice) and Rocky Mountain spotted fever (transmitted by ticks). Beta Proteobacteria Several diverse groups, including Nitrosomonas, which oxidizes ammonia. Pathogenic bacteria in this group include Neisseria gonorrhoeae, which causes gonorrhea. Gamma Proteobacteria Includes the enterobacteria, decomposers that live on decaying plant matter, pathogens, and a variety of bacteria that inhabit humans. Although Escherichia coli is a normal inhabitant of the animal intestinal tract, certain strains can cause moderate to severe diarrhea. One species of Salmonella infects food and produces a toxin that causes a form of food poisoning; another species causes typhoid fever. Vibrios are mainly marine; some are bioluminescent. Vibrio cholerae causes cholera. Pseudomonads are heterotrophs that produce nonphotosynthetic pigments; cause disease in plants and animals, including humans. Purple sulfur bacteria are photoautotrophs that do not produce oxygen. Delta Proteobacteria Includes the myxobacteria (slime bacteria), which secrete slime and glide or creep along. When nutrients are exhausted, these bacteria aggregate into stalked, multicellular reproductive structures called fruiting bodies. Bacterial cells within the fruiting body enter a resting stage. When conditions are favorable, resting cells become active. Epsilon Proteobacteria A small group of bacteria that inhabit the animal digestive tract. Helicobacter can cause peptic ulcers. (continued)

TABLE 25-4

Major Groups of Bacteria (continued) Cyanobacteria contain chlorophyll a and are the only prokaryotes, which, like plants and algae, carry on photosynthesis that generates oxygen. Cyanobacteria were the first organisms that carried on oxygen-generating photosynthesis; considered very important in the evolution of life-forms because their photosynthesis changed the early reducing atmosphere on Earth to an oxidizing atmosphere. Chloroplasts are thought to have evolved from endosymbiotic cyanobacteria. Inhabit ponds, lakes, swimming pools, moist soil, dead logs, tree bark. Some form filaments; other species are solitary. As primary producers, they are an important source of food for marine and freshwater organisms. Some species have special structures that fix nitrogen.

Cengage

Cyanobacteria (Gram-negative)

20 μm

LM of Anabaena, a filamentous cyanobacterium that fixes nitrogen Nitrogen fixation takes place in the rounded cells, called heterocysts.

© David Scharf / Science Faction / Corbis

Gram-positive Bacteria

10 μm

5 μm

© David M. Phillips/ Visuals Unlimited, Inc.

SEM of Actinomycetes naeslundi, a soil-dwelling bacterium that forms filamentous colonies

SEM of Mycoplasma on fibroblast cells

© David M. Phillips/ Visuals Unlimited, Inc.

Chlamydias (Gram-negative)

An extremely diverse group. A few examples follow. Actinomycetes superficially resemble fungi. However, they have peptidoglycan in their cell walls, lack nuclear envelopes, and have other prokaryotic characteristics. Most actinomycetes are saprotrophs that decompose organic materials in soil. Some are anaerobic. Several species of the genus Streptomyces produce antibiotics such as streptomycin, erythromycin, chloramphenicol, and the tetracyclines. Some actinomycetes cause serious lung disease and other infections in humans and other animals. Lactic acid bacteria ferment sugar, producing lactic acid as the main end product. Inhabit decomposing plant material, milk, and other dairy products; responsible for the characteristic taste of yogurt, pickles, sauerkraut, and green olives. Among the normal inhabitants of the human mouth and vagina. Mycobacteria are slender, irregular rods; contain a waxy substance in their cell walls. One species causes tuberculosis; another causes leprosy. Streptococci inhabit the mouth and digestive tract of humans and some other animals. Among the harmful species are those that cause “strep throat,” dental caries, a form of pneumonia, scarlet fever, and rheumatic fever (see chapter opener). Staphylococci normally live in the nose and on skin. Opportunistic pathogens that cause disease when the immunity of the host is lowered. Staphylococcus aureus causes boils and skin infections (some extremely serious); may infect wounds. Certain strains of S. aureus cause a form of food poisoning; other strains cause toxic shock syndrome (see Fig. 25-14). Clostridia are anaerobic. One species causes tetanus; another causes gas gangrene. Clostridium botulinum can cause botulism, an often fatal type of food poisoning. The mycoplasmas are a group of very small bacteria that lack cell walls. They may have evolved from bacteria with gram-positive cell walls. They inhabit soil and sewage; some are parasitic on plants or animals. Some inhabit human mucous membranes but do not generally cause disease; one species causes a mild type of bacterial pneumonia in humans. Chlamydias lack peptidoglycan in their cell walls. They are energy parasites, completely dependent on their hosts for ATP. Infect almost every species of bird and mammal. A strain of Chlamydia causes trachoma, the leading cause of blindness in the world. Sexually transmitted chlamydias are the major cause of pelvic inflammatory disease (PID) in women.

1 μm

TEM of Chlamydia trachomatis in human oviduct cell

© Dennis Kunkel Microscopy / Visuals Unlimited, Inc.

Spirochetes (Gram-negative)

5 μm

LM of Treponema pallidum, the spiro-

Spirochetes are spiral-shaped bacteria with flexible cell walls; move by means of unique internal flagella called axial filaments. Some species are free-living, whereas others form symbiotic associations; a few are parasitic. The spirochete of greatest medical importance is Treponema pallidum, which causes syphilis.

these parasites limit or eradicate males from the population. In some insect species, infected males can reproduce only when they mate with infected females carrying the same Wolbachia strain. In other species, these parasites convert male insects into females. In some wasp species, Wolbachia induces parthenogenesis, development of unfertilized eggs into adult organisms. Because they affect reproduction in their hosts, Wolbachia and other reproductive parasites may influence evolutionary divergence and even extinction in some species. Wolbachia may also affect evolution by horizontal gene transfer.

25.5 IMPACT ON ECOLOGY, TECHNOLOGY, AND COMMERCE ■ ■ LEARNING OBJECTIVES 10 Identify the critical ecological roles played by prokaryotes. 11 Describe some of the important roles played by prokaryotes in commerce and technology.

Many bacteria form biofilms Many types of bacteria that inhabit watery environments form dense films called biofilms that attach to solid surfaces. The bacteria in a biofilm form layers up to 200 mm thick. The bacteria secrete a slimy, gluelike substance rich in polysaccharides and become embedded in this matrix. Biofilms are communities of microorganisms. Most biofilms consist of many species of bacteria and may include other types of organisms, such as archaea, fungi, and protozoa. Scientists have found evidence of ancient biofilms in sediments from coastal marine environments 3.5 billion years old. The dental plaque that forms on teeth is a familiar example of a biofilm (FIG. 25-11). Dental plaque consists of several hundred different types of bacteria and archaea. Biofilms also commonly form on the surfaces of contact lenses and catheters. They sometimes develop on surgical implants such as pacemakers and joint replacements.

Prokaryotes inhabit virtually every environment on Earth and are vital members of the biosphere. They affect other organisms directly and by the ecological roles they play. Prokaryotes produce nitrogen in forms usable by other organisms, and they are a reservoir for nutrients. These microscopic organisms recycle nutrients and are key players in biogeochemical cycles.

Prokaryotes form intimate relationships with other organisms

Some types of bacteria may influence evolution of other species The proteobacteria Wolbachia infect many invertebrates, including insects, spiders, crustaceans, and nematodes (roundworms). Wolbachia are transmitted from generation to generation in the eggs of their hosts, so male hosts are not useful to them. Consequently,

Prokaryotes play key ecological roles Bacteria, especially actinomycetes and myxobacteria, are the most numerous inhabitants of soil. As described earlier in this chapter, many prokaryotes are essential decomposers, chemoheterotrophs

© Steve Gschmeissner /Photo Researchers, Inc.

Prokaryotes interact with other organisms in both beneficial and harmful ways. An intimate relationship between members of two or more species is called symbiosis. The partners in a symbiotic relationship are called symbionts. Symbiotic relationships arise by coevolution. Three forms of symbiosis are mutualism, commensalism, and parasitism. Mutualism is a symbiotic relationship in which both partners benefit. Cows and other ruminants (cud-chewing animals) have mutualistic relationships with bacteria that inhabit their digestive tracts. Ruminants lack enzymes for digesting cellulose. They provide the bacteria with a nutrient-rich home, and the bacteria digest the cellulose for them. Trillions of bacteria inhabit the nutrient-rich human intestine. Some of these are mutualistic bacteria. For example, in exchange for nutrients and a place to live, Bacteroides break down indigestible complex carbohydrates into sugars that their human host can absorb. These bacteria also produce certain vitamins that their host absorbs and uses. In addition, Bacteroides promote proliferation of blood vessels that improve intestinal function. Investigators have reported that these bacteria can turn on specific genes in cells of the host’s intestine; apparently, they can induce synthesis of a compound that may kill competing bacteria. In commensalism, one partner benefits and the other is neither harmed nor helped. Many bacteria that inhabit the human intestine are commensals that live on unused food. In parasitism, one partner lives on or in the other. The parasite benefits and the host is harmed. Disease-causing bacteria are usually not considered obligate parasites because these pathogens typically can survive in other ways.

10 μm

FIGURE 25-11 A familiar biofilm: dental plaque Colorized SEM of dental plaque, which consists of a film of bacteria (red) embedded in a matrix of glycoprotein (blue). Bacteria in the plaque can produce acids that erode tooth enamel, leading to tooth decay.

Prokaryotes are important in many commercial processes and in technology Some microorganisms produce antibiotics. These compounds limit competition for nutrients by inhibiting or destroying other microorganisms. By the 1950s, antibiotics had become important clinical tools that transformed the treatment of infectious disease. Today, about 100 clinically useful antibiotics are available, and literally tons of antibiotics are produced annually. Pharmaceutical companies obtain most antibiotics from three groups of microorganisms: a large group of gram-positive soil bacteria, the actinomycetes; gram-positive bacteria of the genus Bacillus; and molds (eukaryotes belonging to kingdom Fungi). Because of their prolific reproduction rates, bacteria are ideal “factories” for the production of biomolecules. Microbiologists have genetically engineered bacteria to produce certain vaccines, human growth hormone, insulin, and many other clinically im-

Paul Rapson/Science Photo Library

that break down dead organic matter and wastes. They use the products of decomposition as an energy source. When prokaryotes break down organic compounds, many of their components, including nitrogen, oxygen, carbon, phosphorus, sulfur, and certain trace elements, are recycled. (The roles of bacteria in biogeochemical cycles, particularly the nitrogen cycle, are discussed in Chapter 55.) Nitrogen is constantly removed from the soil by plants and other natural processes, as well as by human activities, such as agriculture. Plant growth depends on the availability of usable nitrogen, so it must be continually added to the soil. Several types of bacteria, including cyanobacteria, transform atmospheric nitrogen to forms that can be used by plants (see Table 25-4). Some marine crenarchaeotes also carry on nitrification and may be important in the ocean nitrogen cycle. Rhizobial bacteria, a type of motile bacteria that inhabit the soil, form mutualistic relationships with the roots of legumes, a large family of plants that includes important crops such as beans, peas, and peanuts. The infected plant cells form tumorlike nodules in which the bacteria reside and fix nitrogen (see Fig. 55-9). The bacteria supply the plant with the nitrogen it requires, and the plant provides the bacteria with organic compounds, including sugar needed for cellular respiration. Because many soils are deficient in nitrogen, legumes that have formed mutualistic associations with rhizobial bacteria have a decided advantage over other plants. When the legumes die and are decomposed, the fixed nitrogen is released and enriches the soil. Many prokaryotes, such as the cyanobacteria, carry out photosynthesis, using water as the electron source and generating oxygen. During this process, they fix huge amounts of carbon dioxide into organic molecules. Microbiologists are only beginning to unravel the mysteries of prokaryote ecology. For example, proteobacteria in the SAR11 clade are among the most successful organisms on Earth. Although they are one of the most abundant organisms in the Atlantic Ocean, they were not successfully cultured until 2002, and very little is known about their ecological role. There is some evidence that they are important in cycling carbon, nitrogen, and sulfur in the ocean.

FIGURE 25-12 Bioremediation As they feed on the gasoline and certain other waste products in contaminated soil, certain microorganisms convert hydrocarbons in these pollutants to carbon dioxide and water. Photographed at an oil refinery and chemical plant in the United Kingdom.

portant compounds (see Chapter 15). Researchers are developing genetically engineered bacteria for production of many other medically and agriculturally useful products. Microbial fermentation helps produce many foods and beverages. Lactic acid bacteria are used in producing acidophilus milk, yogurt, pickles, olives, and sauerkraut. Several types of bacteria are used in the production of cheese. Bacteria are involved in making fermented meats such as salami and in producing vinegar, soy sauce, chocolate, certain B vitamins (B12 and riboflavin), and citric acid, a compound added to candy and to most soft drinks. Bacteria are used in bioremediation, the process of using microorganisms (and sometimes other organisms) to detoxify or remove oil, gasoline, and other pollutants or toxic chemicals from the environment. Microorganisms break down certain toxins, leaving behind harmless metabolic byproducts such as carbon dioxide and chlorides (FIG. 25-12). More than 1000 different species of bacteria and fungi have been used to clean up various forms of pollution, and microbiologists are searching for others. For example, bacteria and other microorganisms are used in oil spills to break down oil, oxidizing it to CO2. Microorganisms are also used in sewage treatment and to break down solid wastes in landfills. Archaea are also economically important. Archaea that are adapted to high temperatures or extremely acidic conditions, for example, are a source of enzymes that can be used under these extreme conditions. Archaeal enzymes have been added to laundry and industrial detergents to increase their performance at higher temperatures and pH levels. Another archaeal enzyme has been useful in the food industry to convert cornstarch to dextrins.

Review ■ ■ ■ ■

In what ways do bacteria form relationships with other organisms? What is the composition of a biofilm? What are three ways that prokaryotes are ecologically important? What is bioremediation?

25.6 BACTERIA AND DISEASE ■ ■ LEARNING OBJECTIVES 12 Describe the roles played by Louis Pasteur and Robert Koch in understanding infectious disease; list Koch's postulates.

13 Identify adaptations that have contributed to pathogen success.

Some prokaryote species have coevolved with eukaryotes and are interdependent with them. All plants and animals harbor a population of microorganisms that are considered normal microbiota (also referred to as microflora)—harmless symbiotic prokaryotes. An estimated 700 trillion bacteria normally inhabit the human body! This number greatly exceeds the number of the body’s own cells (about 70 trillion). The presence of certain bacterial populations prevents harmful microorganisms (including other bacteria) from flourishing. A small percentage of bacterial species are important pathogens of plants and animals. Some of the normal bacterial inhabitants are opportunistic pathogens that cause disease only under certain conditions. For example, when the immune system is compromised, opportunistic bacteria increase in number and cause disease. Some important bacterial diseases and the pathogens that cause them are listed in TABLE 25-5. To date, no archaea have been shown to cause specific diseases.

Many adaptations contribute to pathogen success Pathogenic microorganisms can enter the body in food, dust, or droplets or through wounds. Many diseases are transmitted by insect or animal bites. To cause disease, a pathogen must adhere to a specific cell type, multiply, and produce toxic substances. Adherence and multiplication occur only when the pathogen competes successfully with the normal microbiota and counteracts the host’s defenses against invasion. Helicobacter pylori, the most common cause of peptic ulcers (ulcers of the stomach and duodenum), is an extremely successful pathogen (FIG. 25-13). It is also associated with chronic gastritis (stomach inflammation) and with stomach cancer, the second most common type of cancer in the world. Helicobacter pylori inhabits the intestinal tracts of an estimated 40% of adults in developed countries and 80% of adults in developing countries. Among its many adaptations is its ability to produce an alkaline shield around itself that protects it from stomach acid. Also contributing

2 μm

(a) SEM of Helicobacter pylori attached to the epithelial lining of the stomach.

hris Bjornberg/Photo Reachers, Inc.

The idea that some unknown agent caused disease was debated long before Leeuwenhoek discovered microorganisms with his microscope in the late 1600s. However, not until much later did scientists develop the tools or the methods needed to accurately understand the relationships between bacteria and disease. During the late 19th century, several physicians, microbiologists, and chemists working independently laid the foundations for the science of microbiology. French chemist Louis Pasteur disproved the prevailing views of spontaneous generation by demonstrating that sterilization of a sugar and protein culture prevented bacterial growth. Pasteur also developed a rabies vaccine, showing that people can be stimulated to develop immunity to disease. The German physician Robert Koch was the first to clearly demonstrate that bacteria cause infectious disease. In 1876, he showed that Bacillus anthracis caused anthrax. Using a microscope, Koch observed the bacteria in the blood and spleens of dead sheep. When he inoculated mice with the infected sheep blood, he was able to identify B. anthracis in the blood of the mice. He also cultured B. anthracis and showed that when he injected the bacteria into healthy mice they developed anthrax. Koch proposed a set of guidelines, now known as Koch’s postulates, that are still used to demonstrate that a specific pathogen causes specific disease symptoms: (1) the pathogen must be present in every individual with the disease, (2) a sample of the microorganism taken from the diseased host can be grown in pure culture, (3) a sample of the pure culture causes the same disease when injected into a healthy host, and (4) the microorganism can be recovered from the experimentally infected host.

Veronika Burmeister/Visuals Limited

Many scientists have contributed to our understanding of infectious disease

FIGURE 25-13 Helicobacter pylori causes peptic ulcers

TABLE 25-5

Important Bacterial Diseases and Their Causative Agents

Disease

Pathogen

Epidemiology/Comments

Anthrax

Bacillus anthracis

Botulism

Clostridium botulinum

Chlamydia

Chlamydia trachomatis

Cholera

Vibrio cholerae

Diphtheria

Corynebacterium diphtheriae

Epidemic typhus

Rickettsia prowazekii

Gonorrhea Hansen disease (leprosy) Lyme disease

Neisseria gonorrhoeae Mycobacterium leprae Borrelia burgdorferi

Peptic ulcer disease

Helicobacter pylori

Pertussis (whooping cough) Plague

Bordetella pertussis

Pneumonia

Streptococcus pneumoniae Salmonella sp.

Most commonly occurs in domestic animals such as cattle. Can be transmitted to humans from infected animals or animal products. Endospores can live in soil for many years. Anthrax is not spread from person to person. Infection can occur in three forms: cutaneous, by inhalation, and gastrointestinal. Contracted by eating foods that contain the endotoxin or from infected wound. Infant botulism is caused by ingesting endospores. Causes muscle paralysis and can cause death from respiratory failure. One of the most frequently reported sexually transmitted diseases in the U.S. About 75% of infected women and 50% of infected men have no symptoms. If untreated, the infection spreads and damages reproductive organs; can lead to infertility. This pathogen can also infect eyes and causes millions of cases of blindness worldwide each year. Contracted by eating food or drinking water contaminated with the bacterium. Common in areas with inadequate sewage treatment and impure water. Infects intestine and can cause severe diarrhea. Rapid fluid loss can lead to dehydration and death. Transmitted from person to person by intimate respiratory and physical contact. Endemic in developing countries. Not common in U.S. since vaccine became available in 1920s. Infects the heart muscle and respiratory passageways. Transmitted by infected body lice. After an 8- to 12-day incubation period, symptoms include fever, severe headache, muscle aches, and chills. Several days later a rash appears. About 40% of untreated patients die. Common sexually transmitted disease. Thought to be spread from person to person in nasal secretions. Worldwide, this disease has disabled up to 2 million people. Typically affects the skin, nerves, and mucous membranes. Transmitted to humans by bite of infected blacklegged ticks. Symptoms include skin rash, headache, fever, and fatigue. If untreated, infection can spread to joints, heart, and nervous system. Causes peptic ulcer, a lesion in the lining of the stomach or duodenum (upper part of small intestine). Highly communicable from person to person. Causes spasms of severe coughing. Vaccination available. Transmitted from wild rodents, squirrels, and cats to people by infected fleas. If untreated, can cause death. Killed millions of people in Europe during the Middle Ages. Transmitted from person to person. Strains of S. pneumoniae are resistant to one or more antibiotics commonly used to treat this infection. Incidence has decreased since introduction of a vaccine. Transmitted to people in contaminated chicken, eggs, or other food. Also transmitted from feces of infected animals, e.g., lizards, turtles, chicks, birds, even dogs and cats. Symptoms include fever, diarrhea, and stomach pain. Bacteria enter the body through a wound in the skin. Affects the nervous system; causes lockjaw, stiffness in neck, difficulty swallowing, fever, and muscle spasms. Vaccination available. Ingested in contaminated food and water. From 30% to 50% of travelers to high-risk areas (Central and South America, Africa, Middle East, and most of Asia) develop travelers’ diarrhea. The infecting bacteria produce toxins in GI tract. Sexually transmitted disease passed through direct contact with a syphilis sore. If untreated, eventually damages brain, liver, bone, and spleen and can cause death. Transmitted from person to person through inhalation of air containing the pathogen. Symptoms include fatigue, cough, fever, weight loss. Not common in U.S. However, multidrug-resistant form is a growing threat. Transmitted from person to person in food or water contaminated with feces. Risk greatest for travelers in developing countries, but vaccine available. Symptoms include high fever, headache, and loss of appetite. Can cause death if not treated.

Salmonella (salmonellosis)

Yersinia pestis

Tetanus

Clostridium tetani

Travelers’ diarrhea

Escherichia coli is most common cause

Syphilis

Treponema pallidum

Tuberculosis

Mycobacterium tuberculosis

Typhoid fever

Salmonella typhi

to its success are several powerful flagella used to propel the pathogen through the thick mucus lining the stomach. Pathogens produce a variety of substances that increase their success. Some bacteria produce exotoxins, strong poisons that are either secreted from the cell or leak out when the bacterial cell is destroyed. The toxin, not the presence of the bacteria themselves, is responsible for the disease. Diphtheria is caused by a grampositive bacillus (Corynebacterium diphtheriae) that produces a

toxin only when lysogenized by a phage. The diphtheria toxin kills cells and causes inflammation. Botulism, a type of food poisoning that can lead to paralysis and sometimes death, results from eating improperly canned food. Botulism is caused by an exotoxin released by the gram-positive, endospore-forming Clostridium botulinum. During the canning process, food must be heated sufficiently to kill any highly heatresistant endospores that may be present. If not, the endospores

Antibiotic resistance is a major public health problem Bacteria mutate frequently and reproduce rapidly. Many antibiotics target protein synthesis in bacteria. For example, streptomycin and related antibiotics block the initiation of protein synthesis. The tetracyclines block aminoacyl tRNA from binding to the A site on the ribosome. Drug resistance may result from an accumulation of mutations in plasmid or chromosomal DNA. Plasmids that have genes for antibiotic resistance are called R factors. They have genes for resistance to a specific drug and for transferring the resistance to other bacteria. Some R factors have several genes for drug resistance, each encoding resistance to a different drug. Methicillin-resistant Staphylococcus aureus (SA), referred to as MRSA, and vancomycin-resistant Staphylococcus aureus, or VRSA, have been directly linked to the horizontal transfer of antibiotic resistance genes by plasmids during conjugation. An estimated 1 in every 100 healthy people in the United States now carries MRSA (FIG. 25-14). This “superbug” can cause infection in individuals with compromised immune systems. The Centers for Disease Control estimates that MRSA causes more than 90,000 serious infections each year, resulting in more than 17,000 deaths annually. Overuse of antibiotics is the principal contributing factor to drug resistance. In any bacterial population, there are likely at least

■ ■

Janice Haney Carr/CDC

can germinate. The resulting bacterial population grows and releases an exotoxin so powerful that 1 g could kill a million humans! Like many exotoxins, the one that causes botulism is inactivated by heating. (Food must be heated to 80°C for 10 minutes, or boiled for 3 to 4 minutes.) The botulism exotoxin, marketed under the trade name Botox, is used in extremely tiny amounts to treat several medical conditions involving spasms (involuntary muscle contractions). Because Botox is a neurotoxin that works by paralyzing muscles, it can also relax facial wrinkles caused by contraction of the underlying muscles. However, its effects last for only three to eight months. Endotoxins are not secreted by pathogens but are components of the cell walls of most gram-negative bacteria. These compounds affect the host only when they are released from dead bacteria. Endotoxins bind to the host’s macrophages (large phagocytic cells of the immune system) and stimulate them to release substances that cause fever and other symptoms of infection. Unlike exotoxins, which cause specific symptoms, endotoxins appear to affect the entire body. Endotoxins are not destroyed by heating.

1 μm

FIGURE 25-14 Colorized SEM of methicillin resistant Staphylococcus aureus (MRSA). These bacteria also show increased resistance to vancomycin.

a few bacteria that are genetically resistant to a particular antibiotic. The bacteria that are not resistant are killed by the antibiotic, leaving the resistant bacteria to multiply and produce a resistant population. Note that this is a common, present-day example of natural selection. Another type of drug resistance explains why some infections, for example, urinary tract infections, are difficult to cure. When E. coli infect the bladder, the immune system launches a powerful defense. The bacteria subvert the attack by forming biofilms (discussed earlier in this chapter). Each biofilm is surrounded by a matrix rich in polysaccharides and by a protective shell. The bacteria in the biofilm are resistant to antibiotics, as well as to host defenses. By one estimate, bacteria growing in a biofilm are up to 1000 times as resistant to antibiotics as are the same type of bacteria that have not formed a biofilm. Within biofilms, differential gene expression and mutation lead to diverse bacterial colonies. The biofilm strategy also explains some other types of chronic or recurrent infections. A high percentage of infections acquired in hospitals involve biofilms.

Review ■ ■

What important roles do prokaryotes play in ecosystems? Why is each of Koch’s postulates essential?

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

25.1 (page 518) 1 Describe the structure and common shapes of prokaryotic cells. ■

■

Prokaryotic cells are very small and do not have membraneenclosed organelles such as nuclei and mitochondria. Prokaryotic cells have several common shapes: spherical (cocci), rod-shaped (bacilli), and spiral. Spiral bacteria include the spirillum, which is a rigid helix, and spirochete, which is a flexible helix.

2 Contrast the bacterial cell wall in gram-positive and gram-negative bacteria. ■ Most bacteria have cell walls composed of peptidoglycan. The walls of gram-positive bacteria are very thick and consist mainly of peptidoglycan. The cell walls of gram-negative bacteria consist of a thin peptidoglycan layer and an outer membrane resembling the plasma membrane. Some species of bacteria produce a capsule or slime layer that surrounds the cell wall.

Some prokaryotes have hairlike appendages called fimbriae. Pili also extend from the surface of some prokaryotes. Both fimbriae and pili help cells adhere to one another or to certain other surfaces, including cells they infect. 3 Describe movement in prokaryotes, and describe the structure of the bacterial flagellum. Bacterial flagella are structurally different from eukaryotic flagella; ■ each flagellum consists of a basal body, hook, and filament. They produce a rotary motion. ■

Learn more about the structure of prokaryotes and their cell walls by clicking on the figures in CengageNOW.

25.2 (page 521) 4 Describe asexual reproduction in prokaryotes and summarize three mechanisms (transformation, transduction, and conjugation) that may lead to genetic recombination. ■ The genetic material of a bacterium typically consists of a circular DNA molecule and one or more plasmids, smaller circular fragments of DNA. ■ Prokaryotes reproduce asexually by binary fission (the cell divides, forming two cells), budding (a bud forms and separates from the mother cell), or fragmentation (walls form inside the cell, which then separates into several cells). ■ In bacteria, genetic material can be exchanged by transformation, transduction, or conjugation. In transformation, a bacterial cell takes in DNA fragments released by another cell. In transduction, a phage carries bacterial DNA from one bacterial cell into another. In conjugation, a donor bacterium transfers plasmid DNA to a recipient bacterium.

Watch conjugation by clicking on the figure in CengageNOW.

5 State specific factors that contribute to the rapid evolution of bacteria. ■

Rapid reproduction ensures that mutations are rapidly passed to new generations. Horizontal gene transfer—by transformation, transduction, and conjugation—contributes to rapid evolution in bacteria.

■

Some bacteria carry on nitrogen fixation—that is, they reduce nitrogen in the atmosphere to ammonia. Other bacteria convert ammonia to nitrite or nitrate, a process called nitrification.

25.4 (page 525) 7 Compare characteristics of the three domains: Archaea, Bacteria, and Eukarya. Prokaryotes are assigned to domain Archaea and domain Bacteria. ■ Unlike those of bacteria, the cell walls of archaea do not have peptidoglycan. The translational mechanisms of eukaryotes more closely resemble those of archaea than those of bacteria. 8 Distinguish between the two main groups of archaea and identify specific types of archaea belonging to each group. ■ The Crenarchaeota consist mainly of extreme thermophiles, archaea that can inhabit very hot, sometimes acidic, environments. Euryarchaeota include methanogens, extreme halophiles, and some extreme thermophiles. Methanogens are obligate anaerobes that produce methane gas from simple carbon compounds. Extreme halophiles inhabit saturated salt solutions. 9 Describe the main groups of bacteria discussed in this chapter. ■ Major groups of bacteria include proteobacteria (gram-negative), cyanobacteria (gram-negative), gram-positive bacteria, chlamydias (gram-negative), and spirochetes (gram-negative) (see Table 25-4). ■

25.5 (page 530) 10 Identify the critical ecological roles played by prokaryotes. Many bacteria are symbiotic with other organisms. Mutualism is a symbiotic relationship in which both partners benefit. In commensalism, one partner benefits and the other is neither harmed nor helped. In parasitism, the parasite benefits and the host is harmed. Bacterial pathogens cause disease, but are usually not considered obligate parasites. Many bacteria form biofilms, dense communities of bacteria and other microorganisms. ■ Prokaryotes play essential ecological roles as decomposers and are important in recycling nitrogen and other nutrients. Some prokaryotes carry out photosynthesis. 11 Describe some of the important roles played by prokaryotes in commerce and technology. ■ Some bacteria produce antibiotics. We have developed the technology for using certain bacteria to produce vaccines, insulin, and other important compounds. We use bacteria in the production of many foods, including cheese, yogurt, vinegar, and chocolate. We also use microbes in sewage treatment and in bioremediation. ■

25.3 (page 524) 6 Describe the principal modes by which prokaryotes carry on nutrition and energy capture, and compare their requirements for oxygen. ■ Most prokaryotes are heterotrophs that obtain carbon from other organisms; some are autotrophs that make their own organic molecules from simple raw materials. ■ Chemotrophs obtain energy from chemical compounds; phototrophs capture energy from light. ■ Autotrophs may be photoautotrophs, which obtain energy from sunlight, or chemoautotrophs, which obtain energy by oxidizing inorganic chemicals such as ammonia. ■ Photoheterotrophs obtain carbon from other organisms but use chlorophyll and other photosynthetic pigments to trap energy from sunlight. The majority of bacteria are chemoheterotrophs. They are mainly free-living decomposers that obtain both carbon and energy from dead organic matter. ■ Most bacteria are aerobic, that is, they require oxygen for cellular respiration. Some prokaryotes are facultative anaerobes that metabolize anaerobically when necessary; others are obligate anaerobes that can carry on metabolism only anaerobically.

25.6 (page 532) 12 Describe the roles played by Louis Pasteur and Robert Koch in understanding infectious disease; list Koch’s postulates. ■ Louis Pasteur demonstrated that sterilization prevented bacterial growth. Koch’s postulates are a set of guidelines developed by Robert Koch to demonstrate that a specific pathogen causes specific disease symptoms: (1) the pathogen must be present in every individual with the disease, (2) a sample of the microorganism taken from the diseased host can be grown in pure culture, (3) a sample of the pure culture causes the same disease when injected into a healthy host, and (4) the microorganism can be recovered from the experimentally infected host.

13 Identify adaptations that have contributed to pathogen success. ■

Some pathogenic bacteria release strong poisons called exotoxins; others produce endotoxins, poisonous components of their cell walls that are released when bacteria die. Many bacteria have become resistant to antibiotics. Plasmids that have genes for antibiotic resistance are called R factors. See Table 25-5 for a description of some pathogenic bacteria.

T E S T YO U R U N D E R S TA N D I N G 1. Peptidoglycan is a chemical compound found in the cell walls of (a) most viroids (b) most archaea (c) all prokaryotes (d) most bacteria (e) most eukarya 2. Bacterial flagella (a) are homologous with eukaryotic flagella (b) exhibit a rotary motion (c) consist of a basal body and nine pairs of microtubules (d) are important in transduction (e) are characteristic of gram-positive bacteria 3. Endospores (a) are formed by some viruses (b) are extremely durable cells (c) are comparable to the reproductive spores of fungi and plants (d) cause fever and other symptoms in the host (e) are extremely vulnerable to infection by phages 4. In conjugation (a) two bacterial cells of different mating types come together, and genetic material is transferred from one to another (b) a bacterial cell develops a bulge that enlarges and eventually separates from the mother cell (c) fragments of DNA released by a broken cell are taken in by another bacterial cell (d) a phage carries bacterial genes from one bacterial cell into another (e) walls develop in the cell, which then divides into several new cells

hydrogen (b) thermophiles (c) halophiles (d) bacteriophages (e) prokaryotes with cell walls that lack peptidoglycan 9. Rhizobial bacteria (a) are used in the production of yogurt (b) produce antibiotics (c) cause peptic ulcers (d) are the pathogens that cause syphilis (e) form mutualistic relationships with the roots of legumes and fix nitrogen 10. Robert Koch (a) proposed a set of guidelines to demonstrate that a specific pathogen causes specific disease symptoms (b) discovered Helicobacter (c) discovered microorganisms in the late 1600s (d) disproved spontaneous generation (e) demonstrated that people can be stimulated to develop immunity to disease 11. Which group of bacteria contains the gram-positive, anaerobic bacterium that causes botulism? (a) clostridia (b) actinomycetes (c) enterobacteria (d) spirochetes (e) streptococci 12. Label the diagram using Figure 25-2 to check your answers.

5. The majority of heterotrophic bacteria are (a) free-living chemoheterotrophs (b) photoautotrophs (c) chemoautotrophs (d) facultative anaerobes (e) obligate anaerobes 6. Bacteria that are autotrophs (a) do not require atmospheric oxygen for cellular respiration (b) must obtain organic compounds from other organisms (c) manufacture their own organic molecules from simple raw materials (d) get their nourishment from dead organisms (e) produce endospores when oxygen levels are too low for active growth 7. Bacteria that thrive in puncture wounds are likely to be (a) chemoheterotrophs (b) photoautotrophs (c) chemoautotrophs (d) endospores (e) obligate anaerobes 8. Which of the following do not belong to domain Archaea? (a) prokaryotes that produce methane from carbon dioxide and

CRITICAL THINKING 1. Imagine that you discover a new microorganism. After careful study you determine that it should be classified in domain Archaea. What characteristics might lead you to this decision? 2. In what ways would life on Earth be affected if all prokaryotes suddenly became extinct? 3. SCIENCE, TECHNOLOGY, AND SOCIETY. The Centers for Disease Control (CDC) considers antibiotic resistance to be one of the world’s most pressing health problems. What factors are contributing to this problem?

4. EVOLUTION LINK. In what way does the use of antibiotics impose selective pressure on bacteria? Additional questions are available in CengageNOW at www.cengage.com/ login.

26

Protists

Wim van Egmond/Visuals Unlimited, Inc.

Light micrograph (LM) of a marine diatom. The widely distributed diatom Guinardia striata forms colonies composed of spiraling chains of cells.

KEY CONCEPTS

P

26.1 Protists are a diverse group of eukaryotic organisms

trition, and lifestyles. Protists, which include algae, water molds, slime

that vary in body plan (unicellular, colonial, coenocytic, multicellular), method of motility (pseudopodia, cilia, flagella), nutrition type (autotrophic, heterotrophic), and mode of reproduction (asexual, sexual).

molds, and protozoa, are unicellular, colonial, or simple multicellular

rotists are an informal group of primarily aquatic eukaryotic organisms with diverse body forms, types of reproduction, modes of nu-

organisms that have a eukaryotic cell organization (see photograph). The word protist, from the Greek for “the very first,” reflects the idea that

26.2 Much uncertainty surrounds eukaryote evolution, but eukaryote organelles such as chloroplasts probably descended from engulfed cells that survived and became organelles. Current scientific evidence supports splitting the protists and other eukaryotes (land plants, fungi, and animals) into five informal “supergroups.”

protists were the first eukaryotes to evolve.

26.3 Excavates are unicellular protists with atypical,

ever, having a eukaryotic cell structure clearly differentiates protists from

Protists are members of domain Eukarya, the third domain on the tree of life. (Recall the three domains depicted in Figure 23-2.) Eukaryotic cells are characteristic of protists as well as of complex multicellular organisms belonging to the kingdoms Fungi, Animalia, and Plantae. How-

greatly modified mitochondria; they get their name because many have a deep, or excavated, oral groove.

members of the prokaryotic domains Bacteria and Archaea. Recall from

26.4 Chromalveolates are a supergroup of diverse pro-

other membrane-enclosed organelles such as mitochondria and plastids,

tists that may have originated as a result of secondary endosymbiosis in which an ancestral cell engulfed a red alga.

9 + 2 flagella, and multiple chromosomes in which DNA and proteins

26.5 Rhizarians are a supergroup of amoeboid cells that often have tests (shells) through which cytoplasmic projections extend; forams, actinopods, and certain shell-less amoebas are rhizarians.

Chapter 4 that unlike prokaryotic cells, eukaryotic cells have nuclei and

form a complex called chromatin. Sexual reproduction, meiosis, and mitosis are also characteristic of eukaryotes. Until recently, all protists were classified as the kingdom Protista, but this classification was by default: protists included any eukaryotes that

26.6 Archaeplastids include red algae, green algae, and

were not land plants, fungi, or animals. Based largely on recent molecular

land plants, all of which have plastids bounded by outer and inner membranes.

data that have clarified many evolutionary relationships among eukary-

26.7 Unikonts include amoebozoa and choanoflagellates as well as fungi and animals.

otes, kingdom Protista is no longer a recognized clade. Although we have made great progress in understanding the evolutionary relationships among protists, many relationships remain uncertain.

Biologists currently recognize dozens of protist taxa, which, along with other eukaryotes, are classified into five “supergroups.” Consideration of all protist taxa is beyond the scope of this text, but we discuss several representative examples of each supergroup and provide insights into how such diverse eukaryotes may have evolved.

26.1 DIVERSITY IN THE PROTISTS ■ ■ LEARNING OBJECTIVE 1

Discuss in general terms the diversity inherent in protists, including means of locomotion, modes of nutrition, interactions with other organisms, habitats, and modes of reproduction.

Because of their huge numbers, protists are crucial to the natural balance of the living world. Protists are an important source of food for other organisms, and photosynthetic protists supply oxygen to aquatic and terrestrial ecosystems. Certain protists are economically important, and others cause devastating diseases such as malaria. Body plan varies considerably among protists. Most protists are unicellular, with each cell forming a complete organism capable of performing all the functions characteristic of life. Some protists form colonies, loosely connected groups of cells; some are coenocytic, consisting of a multinucleate mass of cytoplasm; and some are multicellular, composed of many cells. Unlike the eukaryotic animals, plants, and many fungi, most multicellular protists have relatively simple body forms without specialized tissues. Size and structural complexity are not the only variable features of protists. During the course of their long evolutionary history, protists have evolved diversity in their means of locomotion, ways of obtaining nutrients, interactions with other organisms, habitats, and modes of reproduction. Protists, most of which are motile at some point in their life cycle, have various means of locomotion. Some move by pushing out cytoplasmic extensions (pseudopodia) along the leading edge

and retracting the cytoplasm that trails behind, as an amoeba does. Other protists move by flexing individual cells; by gliding over surfaces; by waving cilia, short, hairlike organelles; or by lashing flagella, long, whiplike organelles. Some protists have two or more means of locomotion—for example, both flagella and pseudopodia. Methods of obtaining nutrients differ widely among protists. Most algae are autotrophic and photosynthesize as plants do (FIG. 26-1a). Some heterotrophic protists obtain their nutrients by absorption, as fungi do, whereas others resemble animals in that they ingest food. Some protists switch their modes of nutrition and are autotrophic at certain times and heterotrophic at others. Although many protists are free-living, others form stable symbiotic associations with unrelated organisms. These intimate associations range from mutualism, a more or less equal partnership where both partners benefit; to commensalism, where one partner benefits and the other is unaffected; to parasitism, where one partner (the parasite) lives on or in another (the host) and metabolically depends on it (see Chapter 54). Some parasitic protists are important pathogens (disease-causing agents) of plants or animals. Throughout this chapter, we describe specific examples of symbiotic associations involving protists. Most protists are aquatic and live in the ocean or in freshwater streams, lakes, and ponds (FIG. 26-1b). They make up most of the plankton, the floating, often microscopic organisms that inhabit

Amoeba Pseudopodia

Volvox

Scenedesmus Chlamydomonas

Pediastrum

Flagella Flagella

Euglena

Spirogyra

Vorticella

Didinium Cell wall

Flagellum

Cilia

Cilia

Paramecium

Nucleus

Stentor

Chloroplast Cilia Starch granule

(a) Chlamydomonas is a photosynthetic organism with two flagella and a cup-shaped chloroplast.

FIGURE 26-1 Protists

(b) Protists in a drop of pond water. Several modes of locomotion are shown.

surface waters and are the base of the food web in aquatic ecosystems. Other aquatic protists attach to rocks or other surfaces in the water. Even parasitic protists are aquatic because they live in the watery environments of other organisms’ body fluids. Terrestrial protists are restricted to damp places such as soil, cracks in bark, and leaf litter. Reproduction is varied among protists. Almost all protists reproduce asexually, and many also reproduce sexually. However, most protists do not develop multicellular reproductive organs, nor do they form embryos the way more complex organisms do.

Review ■ ■

How do protists vary in their means of obtaining nutrients? What are some of the ways protists interact with other organisms?

26.2 HOW DID EUKARYOTES EVOLVE? ■ ■ LEARNING OBJECTIVES 2 3

Discuss the hypothesis of serial endosymbiosis and briefly explain some of the evidence that supports it. Describe the kinds of data biologists use to classify eukaryotes.

For many years biologists have hypothesized that protists were the first eukaryotic cells and that they evolved from ancestral prokaryotes. However, the more we study eukaryote origins, the more uncertain we are. One thing we can say with absolute certainty is that the evolution of eukaryotes was a complex process. Eukaryotes may have appeared in the fossil record as early as 2.2 billion years ago. Other than a few protists with hard shells, such as diatoms and forams, most ancient protists did not leave many fossils because their bodies were too soft to leave permanent traces. Evolutionary studies of protists focus primarily on molecular and structural comparisons of present-day organisms, which contain many clues about their evolutionary history.

Mitochondria and chloroplasts probably originated from endosymbionts Throughout evolutionary history, one organism has engulfed another to the mutual benefit of both. According to the hypothesis of serial endosymbiosis, certain eukaryotic organelles, particularly mitochondria and chloroplasts, arose from symbiotic relationships between larger cells and smaller bacteria that were incorporated and lived within them. (You might want to review serial endosymbiosis in Figure 21-8.) Cell biologists hypothesize that mitochondria originated from aerobic bacteria. Studies of mitochondrial DNA suggest it is a remnant from the mitochondrion’s past, when it was an independent organism. Ribosomal RNA (rRNA) sequences from mitochondria closely match rRNAs found in purple bacteria, suggesting that ancient purple bacteria were the ancestors of mitochondria. Chloroplast evolution is more complex, given that there were probably several endosymbiotic events (FIG. 26-2). Molecular

evidence supports the view that incorporation of an ancient cyanobacterium within a host cell, known as primary endosymbiosis, resulted in the chloroplasts in today’s red algae, green algae, and land plants. (The host cell was eukaryotic because mitochondria almost certainly evolved before chloroplasts.) Biologists hypothesize that these chloroplasts, which are enclosed by two external membranes, later provided other eukaryotes with their chloroplasts during secondary endosymbiosis. Secondary endosymbiosis occurred frequently in eukaryote evolution, as evidenced by the presence of additional chloroplast membranes. For example, three membranes envelop the chloroplasts of euglenoids and dinoflagellates, and four membranes surround the chloroplasts of diatoms, golden algae, and brown algae. Understanding how these membranes originated is an essential aspect of serial endosymbiosis, and many researchers are studying the origin of chloroplasts in different organisms. Even nonphotosynthetic protists may contain chloroplast relics from secondary endosymbiotic events. Apicomplexans— protists such as Plasmodium, which causes malaria—have a nonfunctional chloroplast surrounded by four membranes. Some researchers hypothesize that drugs that inhibit chloroplasts may also kill Plasmodium. This nonfunctional chloroplast, which is probably derived from a dinoflagellate, may have some medical potential in treating malaria.

A consensus in eukaryote classification is beginning to emerge Scientists re-evaluate evolutionary relationships among the eukaryotes as additional evidence becomes available. Two types of modern research, molecular analysis and ultrastructural studies, have contributed substantially to scientific understanding of the phylogenetic relationships among protists. Molecular data were initially obtained for the gene that codes for small subunit ribosomal RNA in different eukaryotes (SSU rRNA; see Chapter 25). More recently, biologists have compared other nuclear genes, many of which code for proteins, in different protist taxa. Ultrastructure is the fine details of cell structure revealed by electron microscopy. In many cases, ultrastructure data complement molecular data. Electron microscopy reveals similar structural patterns among those protist taxa that comparative molecular evidence suggests are monophyletic—that is, evolved from a common ancestor (see Chapter 23). For example, molecular and ultrastructure data suggest that water molds, diatoms, golden algae, and brown algae—protist taxa that at first glance seem to share few characteristics—are a monophyletic group. Given the diversity in protist ultrastructure and molecular data, biologists regard the protists as a paraphyletic group—that is, protists contain some, but not all, of the descendants of a common eukaryote ancestor. Molecular and ultrastructural analyses continue to help biologists clarify relationships among the various protist phyla and among protists and the other eukaryotic kingdoms. Biologists use these data to develop various classification schemes. A current scheme, which we have adopted in this edition, splits the protists and other eukaryotes (land plants, fungi, and animal) into five informal supergroups (FIG. 26-3 and TABLE 26-1). This

KEY POINT

Scientists hypothesize that plastids have evolved through both primary and secondary endosymbiosis.

Mitochondrion Nucleus Eukaryotic cell with mitochondria Bacterial DNA

(a) Primary endosymbiosis

Eukaryotic cell with mitochondria

(a) In primary endosymbiosis, an ancient eukaryotic cell engulfed a cyanobacterium, which Cyanobacterium survived and evolved into a (ancestor of chloroplast. The ancient cell is chloroplast) depicted as a eukaryotic cell because mitochondria almost certainly evolved before chloroplasts.

(b) In a secondary endosymbiotic event, a heterotrophic eukaryotic cell (with mitochondria) engulfed Chloroplast a eukaryotic cell with chloroplasts DNA (a red alga is depicted). The red (b) Secondary Chloroplast with alga survived and evolved into a two membranes chloroplast surrounded by three endosymbiosis membranes (a dinoflagellate Eukaryotic cell with mitochondria and chloroplasts chloroplast is depicted). Other (red alga) secondary endosymbiotic events resulted in more complex chloroplast membrane structures. Chloroplast with three membranes

Eukaryotic cell with mitochondria and chloroplasts (dinoflagellate)

FIGURE 26-2 Animated Chloroplast evolution by primary and secondary endosymbiosis

classification scheme is different from the eukaryote classification we used in our last edition, and more changes will almost certainly be reported in future editions. Science is an ongoing enterprise, and current knowledge must always be re-evaluated in light of new discoveries. Now that we have a basic understanding of evolution of the protists and other eukaryotes, let’s examine representative protists within the five supergroups.

Review ■ ■

How does serial endosymbiosis explain the origin of chloroplasts? What kinds of scientific evidence support the hypothesis that protists are a paraphyletic group?

26.3 EXCAVATES ■ ■ LEARNING OBJECTIVE 4 Summarize the basic features of excavates and distinguish among diplomonads, parabasilids, and euglenoids.

Excavates are a diverse group of unicellular protists with flagella. These protists are so named because many have a deep, or excavated, oral groove. Unlike other protists, excavates have atypical, greatly modified mitochondria. Many excavates are endosymbionts and live in anoxic (without oxygen) environments. These excavates do not carry out aerobic respiration; they obtain energy by the anaerobic pathway of glycolysis (presumably by fermentation).

KEY POINT

Unraveling phylogenetic relationships among the protists has been difficult, but progress is occurring. Many biologists currently classify eukaryotes into five major groups.

Euglenoids

Excavates

Diplomonads

Dinoflagellates Apicomplexans

Alveolates Water molds

Diatoms

Chromalveolates

Ciliates

Stramenopiles B

Brown algae Golden algae Forams

A Actinopods

Green algae

Land plants

Archaeplastids

Red algae

Rhizarians

Ancestral eukaryote

Amoebas Plasmodial slime molds

Opisthokonts

Fungi

Unikonts

Cellular slime molds

Choanoflagellates Animals

FIGURE 26-3 Evolutionary relationships among eukaryotes This evolutionary tree presents one interpretation of relationships among eukaryotes in which the eukaryotes are placed in five “supergroups.” Organisms within ovals represent separate eukaryotic kingdoms discussed in later chapters. All other eukaryotes are informally known as protists, a paraphyletic group. At node A, all five supergroups

branch from the same point, indicating that the branching order is unresolved at this time. The same is true at node B for the diatoms, brown algae, and golden algae. (Based on Keeling, P. J., “Deep Questions in the Tree of Life,” Science, Vol. 317, Sept. 28, 2007.)

“Protist” Clades

Eukaryote “Supergroup” Excavates Unicellular protists with atypical, greatly modified mitochondria; bikonts

Representative “Protist” Clades

Key Characters

Diplomonads and parabasalids

Two or more flagella; ventral oral (feeding) groove

Euglenoids and trypanosomes

Some with plastids; crystalline rod in flagella

Specific Example © Dr. David Phillips/ Visuals Unlimited, Inc.

TABLE 26-1

Archaeplastids Plastids bounded by outer and inner membranes; include land plants; bikonts

Unikonts Cells that have a single flagellum or are amoebas with no flagella; have a triple-gene fusion that is lacking in other eukaryotes; include animals and fungi

Stramenopiles Water molds, diatoms, brown algae, golden algae

Most have two flagella, one with hairs; no flagella in some

Forams

Porous tests (hard shells) through which cytoplasmic projections (pseudopods) extend

Actinopods

Endoskeletons (internal shells) through which axopods (filamentous pseudopods) extend

Red algae

Chloroplast pigments include phycoerythrin (red pigment) and phycocyanin (blue pigment)

Green algae

Chloroplast pigments identical to those in land plants

Amoebozoa Amoebas, plasmodial slime molds, cellular slime molds

Naked amoebas (no tests) with lobelike pseudopods

Opisthokonts Choanoflagellates

No flagella or single posterior flagellum on motile cells

Currently, excavates include diplomonads, parabasalids, euglenoids, and trypanosomes. The inclusion of these organisms into a single superfamily is somewhat controversial because their relationships to one another are uncertain. Additional studies will be needed to determine if the excavates as currently presented are a monophyletic group.

Diplomonads are small, mostly parasitic flagellates Diplomonads are excavates that have one or two nuclei, no functional mitochondria, no Golgi complex, and up to eight flagella. Giardia is a parasitic diplomonad (FIG. 26-4a). Interestingly, Giardia has two haploid nuclei, each of which contains a complete copy of Giardia’s genome. Giardia lacks functional mitochondria, although it contains certain genes that code for proteins associated with mitochondria in other organisms. Giardia also has reduced

Darkfield LM of Ceratium hirundinella, a dinoflagellate © Wim van Egmond/ Visuals Unlimited, Inc.

Alveoli (flattened vesicles) just inside the plasma membrane

Darkfield LM of Cibicides lobatula, a foram © Wim van Egmond/ Visuals Unlimited, Inc.

Rhizarians Amoeboid cells that often have tests (shells); bikonts

Alveolates Dinoflagellates, ciliates, apicomplexans

Darkfield LM of Micrasterias thomasiana, a green alga © Michael Abbey/ Visuals Unlimited, Inc.

Chromalveolates Diverse protists that may have originated as a result of secondary endosymbiosis in which an ancestral cell engulfed a red alga; bikonts

© Wim van Egmond/ Visuals Unlimited, Inc.

SEM of Trichomonas vaginalis, a parabasalid

Darkfield LM of Amoeba proteus, an amoeba

structures that somewhat resemble mitochondria. This information suggests to some biologists that an early eukaryotic ancestor of Giardia may have possessed mitochondria, which were somehow lost or reduced at a later time during its evolutionary history. Giardia intestinalis is a major cause of water-borne diarrhea throughout the world. Giardia is eliminated as a resistant cyst in the feces of many vertebrate animals. These cysts are a common contaminant in untreated drinking water. In a heavy infection, much of the wall of the small intestine is coated with these flagellates, which interfere with the absorption of digested nutrients and cause weight loss, abdominal cramps, and diarrhea.

Parabasilids are anaerobic endosymbionts that live in animals Parabasilids are anaerobic, flagellated excavates that often live in animals. Trichonymphs and trichomonads are examples of para-

© E. White/Visuals Unlimited, Inc.

Unikonts

Archaeplastids

Rhizarians

Chromalveolates

Excavates

Nuclei

Ancestral eukaryote

2 μm

(a) Diplomonad. This colorized TEM of Giardia intestinalis, a parasitic diplomonad, reveals two nuclei.

The most well-known trichomonad is probably Trichomonas vaginalis, which causes trichomoniasis, a curable sexually transmitted disease (STD) in humans (see SEM in Table 26-1). Trichomoniasis affects both men and women, although the symptoms are more obvious in women. According to the Centers for Disease Control and Prevention, about 7.4 million new cases occur in the United States each year.

© Michael Abbey/Visuals Unlimited, Inc.

Nucleus

Flagella

50 μm

(b) Parabasalid. LM of Trichonympha, a parabasalid that lives in the gut of wood-eating termites and cockroaches. Trichonympha has hundreds of flagella.

FIGURE 26-4 Diplomonads and parabasalids

basilids. Trichonymphs, which have hundreds of flagella, live in the guts of termites and wood-eating cockroaches (FIG. 26-4b). Trichonymphs ingest wood chips from the wood that termites or roaches eat. The trichonymphs rely on endosymbiotic bacteria to digest cellulose in the wood. The insects, trichonymphs, and bacteria all obtain their nutrients from this source. This is an excellent example of mutualism.

Euglenoids and trypanosomes have both free-living species and parasites Euglenoids and trypanosomes are characterized by an unusual flagellum: in addition to the 9 + 2 arrangement of microtubules characteristic of all eukaryotic flagella, these excavates have a crystalline rod in their flagella; the function of this rod is unknown. Like other excavates, euglenoids and trypanosomes also have atypical mitochondria. Most euglenoids are unicellular flagellates, and about onethird of them are photosynthetic (FIGS. 26-5a and b). They generally have two flagella: one long and whiplike and one that is often so short that it does not extend outside the cell. Some euglenoids, such as Euglena, change shape continually as they move through the water because their pellicle, or outer covering, is flexible. Autotrophic euglenoids have the same photosynthetic pigments that green algae and plants have, but euglenoids are not closely related to either group, as shown in Figure 26-3. Some photosynthetic euglenoids lose their chlorophyll when grown in the dark, and they obtain their nutrients heterotrophically, by ingesting organic matter. Other euglenoids are always colorless and heterotrophic. Some heterotrophic species absorb organic compounds from the surrounding water, whereas others engulf bacteria and protists by phagocytosis; they digest the prey within food vacuoles. Trypanosomes are excavates with a single mitochondrion that has an organized deposit of DNA called a kinetoplastid. Trypanosomes are colorless, and many are parasitic and cause disease. In

Flagellum for locomotion

▲

FIGURE 26-5 Euglenoids and trypanosomes

Eyespot

Nonemergent flagellum (indistinguishable in micrograph) Eyespot

Review ■ ■

What cell organelle is atypical in excavates? Give an example of a human disease caused by each of the following: diplomonads, parabasilids, and trypanosomes.

Mitochondria (indistinguishable in micrograph)

Contractile vacuole

Chloroplast

Nucleolus

Nucleus

© Biophoto Associates/Photo Researchers, Inc.

vertebrates, including humans, trypanosomes live in the blood. For example, Trypanosoma brucei is a human parasite that causes African sleeping sickness (FIG. 26-5c). It is transmitted by the bite of infected tsetse flies. Early symptoms include recurring attacks of fever. Later, when the trypanosomes have invaded the central nervous system, infected people are lethargic and have difficulty speaking or walking. At least 50,000 new cases occur each year; if untreated, African sleeping sickness can cause death.

Chromatin

Paramylon body (stored food)

Pellicle

25 μm

(a) LM of Euglena gracilis, a unicellular, flagellate euglenoid.

(b) Euglena’s pellicle is flexible and changes shape easily. The eyespot may shield a light detector at the base of the long flagellum, thereby helping Euglena move to light of an appropriate intensity.

26.4 CHROMALVEOLATES ■ ■ LEARNING OBJECTIVES 6 7

Contrast the two main groups of chromalveolates: alveolates and stramenopiles. Distinguish among the alveolates: dinoflagellates, apicomplexans, and ciliates. Distinguish among the stramenopiles: water molds, diatoms, golden algae, and brown algae.

The chromalveolates are a supergroup composed of extremely diverse protists with few shared characters. Chromalveolates probably originated as a result of secondary endosymbiosis in which an ancestral cell engulfed a red alga (which itself was the result of primary endosymbiosis). Most chromalveolates are photosynthetic, and evidence suggests that heterotrophic chromalveolates, such as the water molds and ciliates, descended from autotrophic ancestors. Classification of the chromalveolates as a monophyletic supergroup is controversial because some DNA sequence data have indicated that they are not monophyletic. The chromalveolates are divided into two main groups, alveolates and stramenopiles The unifying features of protists classified as alveolates include similar ribosomal DNA sequences and alveoli (sing., alveolus), flattened vesicles located just inside the plasma membrane. In some alveolates, the vesicles contain plates of cellulose. Alveolates include the dinoflagellates, apicomplexans, and ciliates. The stramenopiles include water molds, diatoms, golden algae, and brown algae. At first glance, stramenopiles appear too diverse to classify together. However, most stramenopiles have motile cells with two flagella, one of which has tiny hairlike projections extending from the shaft.

Red blood cells

Trypanosome with undulating membrane © Eye of Science/Photo Researchers, Inc.

5

Flagellum

10 μm

(c) SEM of the flagellate Trypanosoma brucei among human red blood cells. Trypanosoma brucei causes African sleeping sickness in humans.

Most dinoflagellates are a part of marine plankton

Unikonts

Archaeplastids

Rhizarians

Chromalveolates

Excavates

Dinoflagellates are generally unicellular, although a few are colonial. Their alveoli contain interlocking cellulose plates impregnated with silicates. The typical dinoflagellate has two flagella. One flagellum wraps around a transverse groove in the center of the cell like a belt, and the other lies in a longitudinal groove (perpendicular to the transverse groove), projecting behind the cell (FIG. 26-6a). The undulation of these flagella propels the dinoflagellate through the water like a spinning top. Indeed, the name is derived from the Greek dinos, meaning “whirling.” Many marine dinoflagellates are bioluminescent.

Many dinoflagellates are photosynthetic (see LM in Table 26-1), but others are heterotrophic and ingest other microorganisms for food. Some dinoflagellates are endosymbionts that live in the bodies of marine invertebrates such as mollusks, jellyfish, and corals (see Fig. 54-12). These symbiotic dinoflagellates, called zooxanthellae, photosynthesize and provide carbohydrates for their invertebrate partners. Zooxanthellae contribute substantially to the productivity of coral reefs. Other dinoflagellates that are endosymbionts lack pigments and are parasites that live off their hosts. Ecologically, dinoflagellates are important producers in marine ecosystems. A few dinoflagellates are known to have occasional population explosions, or blooms. These blooms, known as red tides, frequently color coastal waters orange, red, or brown (FIG. 26-6b). Dinoflagellate blooms are particularly common in warm, nutrient-enriched water. Some dinoflagellate species that form red tides produce a toxin that attacks the nervous systems of fishes, leading to fish kills. Birds sometimes die after eating contaminated fish. Research has also linked red tides to manatee and dolphin deaths in Florida.

Apicomplexans are spore-forming parasites of animals Apicomplexans are a large group of parasitic, spore-forming alveolates, some of which cause serious diseases in humans. They contain the unpigmented remnant of a chloroplast and may have evolved from parasitic dinoflagellates living in the intestines of marine invertebrates. Apicomplexans lack specific structures for locomotion (cilia, flagella, or pseudopodia) and move by flexing.

Ancestral eukaryote

Kevin Schafer /Peter Arnold, Inc. /Alamy

Dr. David Phillips /Visuals Unlimited, Inc.

Flagella

20 μm

(a) SEM of Gymnodinium. Note the two flagella, which are located in grooves.

FIGURE 26-6 Dinoflagellates

(b) Red tide in Mexico. Billions of dinoflagellates produce the orange cloudiness in the water.

Apicomplexans have an apical complex of microtubules that attaches the parasite to its host cell; the apical complex is visible only by using electron microscopy. At some stage in their life cycle, apicomplexans produce sporozoites, small infective agents transmitted to the next host. Many apicomplexans spend part of their complex life cycle in one host species and part in a different host species. Malaria is caused by an apicomplexan (FIG. 26-7). According to the World Health Organization, approximately 250 million people currently have malaria, and about 800,000 people, mostly children in developing countries, die from the disease each year. Although for centuries Chinese, Greek, Arabic, and Roman writings described the disease, its causative agent and mode of transmission were not identified until the end of the 19th century. British sci-

KEY POINT

entist Ronald Ross received the Nobel Prize in 1902 for his role in elucidating the life cycle of Plasmodium, the apicomplexan that causes malaria. Today malaria is recurring in many countries where it was under control for decades. Formerly effective control methods— antimalarial drugs and pesticides—have lost much of their effectiveness. Chloroquine and several other antimalarial drugs are taken prophylactically to prevent malaria, but Plasmodium has evolved resistance to several of these. Moreover, mosquitoes have evolved resistance to many pesticides. Researchers are currently testing new antimalarial drugs and several possible vaccines against malaria. The sequencing of the genomes of P. falciparum and the Anopheles mosquito may lead to new diagnostics, drugs, and vaccines.

Plasmodium, an apicomplexan that is the causative agent of malaria, lives in two hosts, mosquitoes and humans.

1

Infected female Anopheles mosquito bites uninfected human and transmits Plasmodium sporozoites to human blood. Anopheles mosquito

Liver

2

6

Merozoites released

Zygote embeds in mosquito’s stomach lining and produces sporozoites (spores), which are released and migrate to salivary glands.

DIPLOID (2n) Zygote (2n)

Fertilization

HAPLOID (n)

Sporozoites enter liver cells and divide to produce merozoites. Merozoites released from liver cells infect red blood cells.

Red blood cells 3

Anopheles mosquito Gametes

In blood cells, merozoites divide to form more merozoites, which infect more red blood cells. Some merozoites form gametocytes. © Dr. Gopal Murti/Photo Researchers, Inc.

Meiosis

Liver cell

Sporozoites (n)

Gametocytes 5

In mosquito’s digestive tract, gametocytes develop into gametes, and fertilization occurs.

4

Uninfected female Anopheles mosquito bites infected person and obtains Plasmodium gametocytes. Merozoites 3 μm Red blood cell

FIGURE 26-7 Animated Life cycle of Plasmodium, the causative agent of malaria The inset shows a TEM of a colorized human red blood cell filled with merozoites.

Not all ciliates are motile. Some sessile forms have stalks, and others, although capable of some swimming, are more likely to remain attached to a rock or other surface at one spot. Their cilia set up water currents that draw food toward them. Ciliates differ from other protists in having two kinds of nuclei: one or more small, diploid micronuclei that function in reproduction; and a larger, polyploid macronucleus that controls cell metabolism and growth. Most ciliates are capable of a sexual process called conjugation, in which two individuals come together and

Ciliates use cilia for locomotion Ciliates are among the most complex of eukaryotic cells. These unicellular alveolates have a pellicle that gives them a definite but changeable shape. In Paramecium, the surface of the cell is covered with several thousand fine, short, hairlike cilia that extend through pores in the pellicle to facilitate movement (FIGS. 26-8a and b; also see Fig. 4-4c). The cilia beat with such precise coordination that the organism can back up and turn around as well as move forward.

Cilia Food vacuoles Food Micr onucleus

© Robert Brons / Biologcal Photo Service

Macronucleus Contractile vacuole

Anal pore

Oral groove Contractile vacuole

Food vacuole

50 μm

(a) Note the complex cell structure in this phase contrast LM of Paramecium, a freshwater ciliate. Like many ciliates, Paramecium has multiple nuclei: a macronucleus and one or more smaller micronuclei. Disintegrating macronuclei

Macronuclei

2 First meiotic Two sexually division in compatible each cell individuals join at oral surfaces

3

Second meiotic division in each cell

(c) Conjugation in Paramecium caudatum.

FIGURE 26-8 Ciliates

Diploid nuclei (2n)

Disintegrating micronuclei

Micronuclei (2n) 1

(b) Food particles are swept into Paramecium’s ciliated oral groove and incorporated into food vacuoles. Lysosomes fuse with the food vacuoles, and the food is digested and absorbed; undigested wastes are eliminated through the anal pore.

4

One haploid 5 Each conjugating 6 Haploid micronucleus cell exchanges a micronuclei fuse divides by micronucleus mitosis; others disintegrate

7

Cells separate

exchange genetic material (FIG. 26-8c). Conjugation results in two “new” cells that are genetically identical to each other but different from what they were before conjugation. Mitosis and cell division need not follow immediately after conjugation. Ciliates usually divide perpendicularly to their longitudinal axis.

Water molds produce biflagellate reproductive cells Water molds were once classified as fungi because of their superficial resemblance. Both water molds and fungi have a body, called a mycelium, that grows over organic material, digesting it and then absorbing the predigested nutrients (FIG. 26-9). The thread-

like hyphae that make up the mycelium in water molds are coenocytic, meaning that there are no cross walls, and the body of these stramenopiles consists of a single multinucleate cell. The cell walls of water molds are composed of cellulose (as in plants), chitin (as in fungi), or both. When food is plentiful and environmental conditions are favorable, water molds reproduce asexually. A hyphal tip swells and a cross wall is formed, separating the hyphal tip from the rest of the mycelium. Within this structure, called a zoosporangium, tiny biflagellate zoospores form, each of which swims about, lands and encysts, and eventually develops into a new mycelium. When environmental conditions worsen, water molds initiate sexual reproduction. After fusion of male and female nuclei, thick-walled Oosphere nucleus Haploid sperm nucleus

2

Antheridium (male reproductive structure)

1

Meiosis results in haploid sperm nuclei within antheridia and haploid oospheres (eggs) within oogonia.

3

Meiosis

Sperm nuclei move into oospheres.

Fertilization HAPLOID (n) GENERATION

Saprolegnia reproduces sexually by antheridia and oogonia.

Oospores

After fertilization, oospores develop from fertilized oospheres. Each oospore may develop into new mycelium.

4

SEXUAL REPRODUCTION

Oogonium (female reproductive structure)

DIPLOID (2n) GENERATION

Germination of oospore

Mycelium Germination of the zoospore

© James Richardson/Visuals Unlimited, Inc.

Encysted secondary zoospore Secondary zoospore (bean-shaped)

Zoosporangium

ASEXUAL REPRODUCTION (by mitosis) 5

Encysted primary zoospore Mycelium on dead insect.

Zoospores

Primary zoospore (pear-shaped)

FIGURE 26-9 Water molds The life cycle of Saprolegnia, a water mold. Inset: A mycelium of Saprolegnia radiates from a dead insect.

Saprolegnia reproduces asexually by forming zoospores within zoosporangium.

© Dr. Richard Kessel & Dr. Gene Shih/Visuals Unlimited, Inc.

oospores develop from the oospheres (female gametes). Water molds often spend the winter as oospores. Some water molds have played a profound role in human history. For example, the Irish potato famine of the 19th century was precipitated by the water mold Phytophthora infestans, which causes late blight of potatoes. (The genus Phytophthora is named from Greek words meaning “plant destruction.”) During several rainy, cool summers in Ireland in the 1840s, the water mold multiplied unchecked, causing potato tubers to rot in the fields. Because potatoes were the staple of the Irish peasants’ diet, as many as one million people starved. The famine prompted a mass migration out of Ireland to the United States and other countries. A close relative of the late blight water mold, P. ramorum, causes sudden oak death, which is killing oak forests in several western states. Plant pathologists are concerned that the disease may spread to Midwestern and Eastern forests. This particular water mold also attacks redwoods, Douglas firs, bay trees, maples, and several other plant species, but most have only twig and leaf infections, not the rapid death observed in oaks.

20 μm

(a) SEM of a unicellular diatom. Note the striking shell that contains silica.

Diatoms have shells composed of two parts Most diatoms are unicellular stramenopiles, although a few exist as colonies (see chapter opener figure). The cell wall of each diatom consists of two shells that overlap where (b) Asexual reproduction in diatoms. After cell division, each new cell they fit together, much like a petri dish. Silica is deposited retains half of the original shell. The newly synthesized half of the shell in the shell, and this glasslike material is laid down in inalways fits inside the original half. As a result, one of the new cells is slightly tricate patterns (FIG. 26-10a). There are two basic groups smaller. of diatoms: those with radial symmetry (wheel shaped) and those with bilateral symmetry (boat shaped or needle FIGURE 26-10 Diatoms shaped). Although some diatoms are part of the floating plankton, others live on rocks and sediments, where they filtering agent, diatomaceous earth is used to refine raw sugar and move by gliding. This gliding movement is facilitated by the secreto process vegetable oils. Because of its abrasive properties, diatotion of a slimy material from a small groove along the shell. maceous earth is a common ingredient in scouring powders and Diatoms most often reproduce asexually by mitosis. When a metal polishes; it is no longer added to most toothpastes because diatom divides, the two halves of its shell separate, and each beit is too abrasive for tooth enamel. The intricately detailed diatom comes the larger half of a new diatom shell (FIG. 26-10b). Because the glass shell cannot grow, some diatom cells get progressively shells are often used to test microscope resolution down to 1 mm. smaller with each succeeding generation. When a diatom reaches a fraction of its original size, sexual reproduction occurs, with the Brown algae are multicellular stramenopiles production of shell-less gametes. Sexual reproduction restores the Brown algae are the largest and most complex of all algae comdiatom to its original size because the resulting zygote, a 2n cell monly called seaweeds. All brown algae are multicellular and range that results from the fusion of n gametes, grows substantially bein size from a few centimeters (about an inch) to 75 m (about fore producing a new shell. 260 ft). Their body forms are branched filaments; tufts; fleshy Diatoms are common in fresh water but are especially abun“ropes”; or thick, flattened branches. The largest brown algae, dant in relatively cool ocean water. They are ecologically significant called kelps, are tough and leathery in appearance. as major producers in aquatic ecosystems. At least one species is Many kelps have leaflike blades in which most photosyntoxic and linked to shellfish poisonings, marine mammal strandthesis occurs, stemlike stipes, and rootlike anchoring holdfasts ings, and the deaths of sea lions along the central California coast. (FIG. 26-11a). They often have gas-filled bladders that provide buoyWhen diatoms die, their shells trickle to the ocean floor and acancy. (The blades, stipes, and holdfasts of brown algae are not cumulate in layers that eventually become sedimentary rock. After homologous to the leaves, stems, and roots of plants. Brown algae millions of years, geologic upheaval has exposed some of these deand plants arose from different unicellular ancestors, as shown in posits on land. Called diatomaceous earth, these deposits are mined Figure 26-3.) and used as filtering, insulating, and soundproofing materials. As a

© Gregory Ochocki/Photo Researchers, Inc.

Stipe Holdfast

(a) Laminaria is widely distributed on rocky coastlines of temperate and polar seas. It grows to 2 m (6.5 ft).

FIGURE 26-11 Brown algae

(b) A kelp (Macrocystis pyrifera) bed is ecologically important to aquatic organisms, including the sea lion shown here. Photographed off the coast of California.

© Philip Sze/Visuals Unlimited, Inc.

Reproduction is varied and complex in the brown algae. Their Most golden algae are unicellular biflagellates reproductive cells, both asexual zoospores and sexual gametes, are Golden algae are found in both freshwater and marine enviusually biflagellate. Most have a life cycle that exhibits alternation ronments. Most species are biflagellate, unicellular organisms, of generations, in which they spend part of their life as multicelalthough some are colonial (FIG. 26-12a). A few of these stramenolular haploid organisms and part as multicellular diploid organisms piles lack flagella and are similar to amoebas in appearance except (see Fig. 10-19c). that golden algae contain chloroplasts. Tiny scales of either silica Brown algae are commercially important for several reasons. Their cell walls contain a polysaccharide called algin that is harvested from kelps such as Macrocystis and used as a thickening and stabilizing agent in ice cream, toothpaste, shaving cream, hair spray, and hand lotion. Brown algae are an important human food, particularly in eastern Asia, and they are rich sources of certain vitamins and minerals such as iodine. Brown algae are common in cooler marine waters, especially along rocky coastlines, where they live mainly in the intertidal zone or relatively shallow offshore waters. Kelps form extensive underwater “forests,” or kelp beds (FIG. 26-11b). They are essential in that ecosystem as im1 μm 10 μm portant food producers, and they provide habitat for many marine invertebrates, (a) LM of a colonial golden alga (Synura) (b) SEM of a coccolithophorid (Emiliania found in freshwater lakes and ponds. fishes, and mammals. The diversity of life huxleyi ). Note the overlapping scales of supported by kelp beds rivals that found calcium carbonate. FIGURE 26-12 Golden algae in coral reefs.

Dr. Elizabeth Venrick/Scripps Institution of Oceanography

© J. R. Waaland/Biological Photo Service

Blade

Unikonts

Archaeplastids

Rhizarians

Chromalveolates

Excavates

or calcium carbonate may cover the cells. Reproduction in golden algae is primarily asexual and involves the production of biflagellate, motile spores called zoospores. Most golden algae are photosynthetic, and the pigment composition of golden algae gives them a golden or golden brown color. A few species ingest bacteria and other particles of food. Ecologically, golden algae are important producers in marine environments. They compose a significant portion of the ocean’s nanoplankton, extremely minute algae (2 mm to 10 mm) that are major producers because of their great abundance. Classification of golden algae is controversial. Some biologists lump diatoms and golden algae in a single phylum, whereas others classify both groups as brown algae. At the other extreme, some biologists divide the golden algae into two phyla by placing many of the marine species, such as coccolithophorids (FIG. 26-12b), in a separate phylum.

Review ■

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■ ■

What are the two main groups of chromalveolates? What are their distinguishing characters? Why do some biologists think the apicomplexans descended from dinoflagellates? Which water mold has influenced human history? Explain your answer. What is the ecological significance of the diatoms? the brown algae?

26.5 RHIZARIANS ■ ■ LEARNING OBJECTIVE 8

Describe the forams and actinopods, and explain why many biologists classify them in the rhizarian supergroup.

Rhizarians are a diverse supergroup of amoeboid cells that often have hard outer shells, called tests, through which cytoplasmic projections extend. The threadlike cytoplasmic projections suggest the name rhizarian, from the Greek rhiza, meaning “root.” Forams and actinopods are rhizarians, as are certain shell-less amoebas. Not all amoebas are rhizarians, however, and many amoeba species are more closely related to other eukaryotic clades. Current molecular evidence indicates that the rhizarian supergroup is monophyletic.

Forams extend cytoplasmic projections that form a threadlike, interconnected net Almost all foraminiferans (forams) are marine rhizarians that produce elaborate tests (FIG. 26-13a; also see LM in Table 26-1). The ocean contains enormous numbers of forams, which secrete chalky, many-chambered tests with pores through which cytoplasmic projections are extended. (Foraminifera is derived from the Latin for “bearing openings.”) The cytoplasmic projections form a sticky, interconnected net that entangles prey. Many forams contain unicellular algal endosymbionts (green algae, red algae, or diatoms)

Biophoto Associates

© Robert Brons / Biological Photo Service

Ancestral eukaryote

500 μm

(a) SEM of a foram test. Note the pores through which cytoplasm extrudes.

FIGURE 26-13 Rhizarians

100 μm

(b) LM of an unidentified living actinopod from the Red Sea. Many slender axopods project from the cell. The shell is not visible because it is an endoskeleton and cytoplasm covers it on all sides.

that provide food by photosynthesis. Many foram species live on the ocean floor, but others are part of the plankton. Dead forams settle on the bottom of the ocean, where their tests form a gray mud that is gradually transformed into chalk. With geologic uplifting, these chalk formations become part of the land, for example, the White Cliffs of Dover in England. (The White Cliffs of Dover are the remains of a variety of carbonate organisms, not only forams.) Because foram tests often appear in rock layers covering oil deposits, geologists exploring for oil look for foram tests in rock strata. Forams are well preserved in the fossil record, and biologists use some as index fossils, markers to help identify ancient sedimentary rock layers (see Chapter 18).

prey inside the body. Many actinopods contain algal endosymbionts that provide them with the products of photosynthesis. Some actinopods, called radiolarians, secrete elaborate, beautiful glassy shells made of silica. Radiolarians are an important constituent of marine plankton. When radiolarians and other actinopods die, their shells settle and become an ooze (sediment) that may be several meters thick on the ocean floor.

Review ■ ■

Why are forams and actinopods classified as rhizarians? What character does the term rhizarian indicate about the forams and actinopods?

Actinopods project slender axopods

Unikonts

Archaeplastids

Rhizarians

Chromalveolates

Excavates

Actinopods are mostly marine plankton rhizarians with long, filamentous cytoplasmic projections called axopods that protrude through pores in their shells (FIG. 26-13b). A cluster of microtubules strengthens each axopod. Unicellular algae and other prey become entangled in these axopods and are engulfed outside the main body of the actinopod; cytoplasmic streaming carries the

26.6 ARCHAEPLASTIDS ■ ■ LEARNING OBJECTIVE 9 Describe evidence supporting the hypothesis that red algae and green algae should be included in a monophyletic group with land plants.

In the classification scheme adopted in this text, the monophyletic group of archaeplastids includes red algae and green algae, which are discussed here, and land plants, which are in a separate kingdom (see Chapters 27 and 28). Biologists classify these groups together based on molecular data and on the presence of chloroplasts bounded by outer and inner membranes, suggesting that they developed directly from a cyanobacterial endosymbiont. All photosynthetic protists other than archaeplastids have plastids surrounded by three or four membranes.

Red algae do not produce motile cells The vast majority of red algae are multicellular organisms, although there are a few unicellular species. The multicellular body form of red algae commonly consists of complex, interwoven filaments that are delicate and feathery (FIG. 26-14a); a few red algae

(a) Polysiphonia, which is widely distributed throughout the world, has a highly branched body of interwoven filaments.

FIGURE 26-14 Animated Red algae

© foodfolio /Alamy

© Philip Sze / Visuals Unlimited, Inc.

© D. Gotshall /Visuals Unlimited, Inc.

Ancestral eukaryote

(b) Bossiella is a coralline red alga encrusted with calcium carbonate. It lives in the Pacific Ocean.

(c) Nori, dried and pressed sheets of the red alga Porphyra, surrounds sushi.

© Tom Adams / Visuals Unlimited, Inc.

are flattened sheets of cells. Most multicellular red algae attach to rocks or other substrates by a basal holdfast. Reproduction in the red algae is remarkably complex, with an alternation of sexual and asexual stages. No flagellate cells develop during the life cycle. Red algae primarily live in warm tropical ocean waters, although a few species occur in fresh water and in soil. Some red algae, known as coralline algae, incorporate calcium carbonate in their cell walls from the ocean water (FIG. 26-14b). The hard calcium carbonate may protect coralline algae from the rigors of wave action. These coralline red algae build “coral” reefs and are perhaps as crucial as coral animals in this process. The cell walls of red algae often contain thick, sticky polysaccharides that have commercial value. For example, agar is a polysaccharide extracted from certain red algae used as a food thickener and culture medium, a substrate on which to grow microorganisms and propagate some plants, such as orchids. Another poly-

50 μm

Green algae share many similarities with land plants Green algae have pigments, energy reserve products, and cell walls that are chemically identical to those of land plants. Green algae are photosynthetic, with chloroplasts of a wide variety of shapes. Most green algae have cell walls with cellulose, although some lack walls. Because of these and other similarities, biologists generally accept that land plants arose from ancestral green algae (see Fig. 26-3). Based on recent molecular and ultrastructure data, some biologists classify this diverse group in the plant kingdom. Green algae exhibit a variety of body types, from single cells to colonial forms, to coenocytic algae (multinucleate), to multicellular filaments and sheets (FIG. 26-15; also see LM in Table 26-1). The multicellular forms do not have cells differentiated into tissues, a characteristic that separates them from land plants. Most green algae have, or produce, flagellate cells during their life cycle, although a few are totally nonmotile. Reproduction in the green algae is as varied as their body forms, with both sexual and asexual reproduction. Many green algae have life cycles with an alternation of haploid and diploid generations. Asexual reproduction is by mitosis and cell division in single cells or by fragmentation in multicellular forms. Many green algae produce spores asexually by mitosis; if these spores have flagella and are motile, they are called zoospores (FIG. 26-16). Sexual reproduction in the green algae involves gamete formation

100 μm

(b) LM of two Volvox colonies, each composed of up to 50,000 cells. New colonies are inside the parental colonies, which eventually break apart.

FIGURE 26-15 Green algae

James W. Perry

© J. Robert Waaland/Biological Photo Service

J. M. Kingsbury in memory of William T. Keeton

(a) LM of Spirogyra, a filamentous green alga with spiral chloroplasts.

saccharide extracted from red algae, carrageenan, is a food additive used to stabilize chocolate milk and to provide a thick, creamy texture to ice cream and other soft processed foods. Carrageenan is also used to stabilize paints and cosmetics. Red algae are a source of vitamins (particularly A and C) and minerals, especially in Japan and other eastern Asian countries where people eat red algae fresh, dried, or toasted in such traditional foods as sushi and nori (FIG. 26-14c).

(c) Ulva’s thin, sheetlike form suggests its common name, sea lettuce.

(d) Chara, a green alga commonly called a stonewort, is closely related to land plants. Chara is widely distributed in fresh water.

5

– – – ASEXUAL REPRODUCTION (by mitosis)

Both mating types reproduce asexually by mitosis; only (−) strain is shown.

Zoospores

–

– 4

Four haploid cells emerge, two (+) and two (−).

1

+ –

–

–

Gametes are produced by mitosis.

SEXUAL REPRODUCTION

+

–

HAPLOID (n) GENERATION

+

–

from a different strain

DIPLOID (2n) GENERATION

+ Meiosis

3

Fertilization

2

(+) and (−) gametes fuse, forming a diploid zygote.

Meiosis occurs. Zygote (2n)

FIGURE 26-16 Animated The life cycle of Chlamydomonas Chlamydomonas is a unicellular, haploid green alga with two mating types, (+) and (−).

in unicellular gametangia (sing., gametangium), reproductive structures in which gametes are produced. Green algae are found in both aquatic and terrestrial environments. Aquatic green algae primarily inhabit fresh water, although there are many marine species. Terrestrial green algae are restricted to damp soil, cracks in tree bark, and other moist places. Many green algae are symbionts with other organisms; some live as endosymbionts in body cells of invertebrates, and a few grow together with fungi as “dual organisms” called lichens (discussed in Chapter 29). Regardless of where they live, green algae are ecologically important as producers.

Review ■

Why do many biologists classify red algae and green algae with land plants?

26.7 UNIKONTS ■ ■ LEARNING OBJECTIVE 10 Briefly describe and compare the following unikonts: amoebas, plasmodial slime molds, cellular slime molds, and choanoflagellates.

Unikonts are a supergroup composed of certain amoebas, plasmodial slime molds, cellular slime molds, choanoflagellates, fungi (discussed in Chapter 29) and animals (discussed in Chapters 30 through 32). Unikonts share a single posterior flagellum in flagellate cells such as sperm and motile spores, although some extant organisms in this supergroup have lost the flagellum. Many unikonts also have a single centriole. Examine Figure 26-3 again and note that the unikonts are divided into two clades. One of

these is the opisthokonts, which consist of fungi, choanoflagellates, and animals. You will learn more about the opisthokonts in Chapters 29 and 30. Unikonts also have a triple-gene fusion that has major evolutionary significance. In a triple-gene fusion, three separate genes fused into a single unit early in the course of eukaryote evolution; the fused gene codes for a multienzyme protein. You may recall that Figure 26-3 showed all five supergroups branching from the same point, indicating that the branching order is unresolved. However, triple-gene fusion provides evidence of a bifurcation, or branch of the eukaryotes into two main clades, close to the root of the common ancestor of all eukaryotes (FIG. 26-17). The two branches consist of (1) unikonts, which had a common ancestor with a single posterior flagellum, and (2) all other eukaryotes, collectively called bikonts, which had a common ancestor with two flagella.

KEY EXPERIMENT QUESTION: How do we deduce details of eukaryote evolution near the root of the evolutionary tree? HYPOTHESIS: Derived gene fusions, which have occurred rarely during the course of evolution, can provide insights into eukaryote evolutionary relationships. EXPERIMENT: Alexandra Stechmann and Thomas Cavalier-Smith of Oxford University sequenced a double-gene fusion that codes for two enzymes, dihydrofolate reductase (DHFR) and thymidylate synthase (TS). In a second experiment, they sequenced a triple-fused gene involved in the enzymatic synthesis of pyrimidine nucleotides. They tested for the presence of these two gene fusions in various eukaryote groups as well as in bacteria and archaea. Bikonts

Excavates Chromalveolates Rhizarians Archaeplastids

Unikonts

Double-gene fusion (DHFR-TS)

Triple-gene fusion (pyrimidine nucleotide synthesis) Ancestral eukaryote

RESULTS AND CONCLUSION: The double-gene fusion is present in all eukaryotes examined except for unikonts; bacteria and archaea also lack the double-gene fusion. The unikonts have the triple-gene fusion that is lacking in all other eukaryotes, bacteria, and archaea. These gene fusions suggest a fundamental division of eukaryotes into two main clades—the unikonts and all other eukaryotes, collectively called bikonts. Source: Stechmann, A. and T. Cavalier-Smith. “The Root of the Eukaryote Tree Pinpointed.” Current Biology, Vol. 13, Sept. 2, 2003.

FIGURE 26-17 Stechmann and Cavalier-Smith’s research on evolution of the protists

Amoebozoa are unikonts with lobose pseudopodia Most amoebozoa produce temporary cytoplasmic projections called pseudopodia (sing., pseudopodium, meaning “false foot”) at some point in their life cycle. The pseudopodia of amoebozoa are lobose—that is, rounded and wide—as opposed to the slender cytoplasmic projections characteristic of rhizarians. Biologists currently classify amoebas, plasmodial slime molds, and cellular slime molds as amoebozoa. Amoebas move by forming pseudopodia Amoebas are unicellular amoebozoa found in soil, fresh water, the ocean, and other organisms (as parasites). Because of the extreme flexibility of their outer plasma membrane, many members of this group have an asymmetrical body form and continually change shape as they move. (The word amoeba derives from a Greek word meaning “change.”) An amoeba moves by pushing out lobose pseudopodia from the surface of the cell (see LM in Table 26-1). More cytoplasm flows into the pseudopodia, enlarging them until all the cytoplasm has entered and the organism as a whole has moved. Pseudopodia also capture and engulf food by surrounding and forming a vacuole around it (FIG. 26-18). Food particles are digested when the food vacuole fuses with a lysosome containing digestive enzymes. Digested materials are absorbed from the food vacuole, which gradually shrinks as it empties. Amoebas reproduce asexually, splitting into two equal parts after mitotic division of the nucleus; sexual reproduction has not been observed. Parasitic amoebas include Entamoeba histolytica, which causes amoebic dysentery, a serious human intestinal disease characterized by severe diarrhea, bloody stools, and ulcers in the intestinal wall. In especially severe cases, the organism spreads from the large intestine and causes abscesses in the liver, lungs, or brain. Entamoeba histolytica is transmitted as cysts in contaminated drinking water. A cyst is a thick-walled, resistant, resting stage in the life cycle of some protists. Other amoebas, such as Acanthamoeba, are usually free-living but produce opportunistic infections such as eye infections in contact lens wearers. Plasmodial slime molds feed as multinucleate plasmodia The feeding stage of a plasmodial slime mold is a plasmodium, a multinucleate mass of cytoplasm that can grow up to 30 cm (1 ft) in diameter (FIG. 26-19a). The slimy plasmodium streams over damp, decaying logs and leaf litter, often forming a network of channels that covers a large surface area. As it creeps along, it ingests bacteria, yeasts, spores, and decaying organic matter. When the food supply dwindles or there is insufficient moisture, the plasmodium crawls to an exposed surface and starts reproducing. Stalked structures of intricate complexity and beauty usually form from the drying plasmodium (FIG. 26-19b). Within these structures, called sporangia, meiosis produces haploid spores that are extremely resistant to adverse environmental conditions. When conditions become favorable, the spores germinate, and a haploid reproductive cell emerges from each. This haploid cell is either a biflagellate swarm cell or an amoeboid myxamoeba,

Unikonts

© Michael Abbey/ Photo Researchers, Inc.

Archaeplastids

Rhizarians

Chromalveolates

Excavates

Ancestral eukaryote

Green alga

Pseudopodia 100 μm

FIGURE 26-18 Animated Amoeba LM of a giant amoeba (Chaos carolinense). This unicellular protist, which moves and feeds by pseudopodia, is surrounding and ingesting a colonial green alga. Chaos amoebas are generally scavengers that feed on debris in freshwater habitats, but they ingest living organisms when the opportunity arises.

depending on available moisture; flagellate cells form in wet conditions. Swarm cells and myxamoebas act as gametes, which fuse to form a zygote with a diploid nucleus. The resultant diploid nucleus divides many times by mitosis, but the cytoplasm does not divide, so the result is a multinucleate plasmodium. The plasmodial slime mold Physarum polycephalum is a model organism that researchers use to study many fundamental biological processes, such as growth, cytoplasmic streaming, and the function of the cytoskeleton.

R. Calentine/CBR Images

R. Calentine/CBR Images

Cellular slime molds feed as individual amoeboid cells The cellular slime molds are amoebozoa with close affinities to amoebas and plasmodial slime molds. During its feeding stage, each cellular slime mold is an individual amoeboid cell that behaves as a separate, solitary organism (FIG. 26-20). Each cell creeps over rotting logs and soil or swims in fresh water, ingesting bacteria and other particles of food as it goes. Each amoeboid cell has a haploid nucleus and reproduces by mitosis, as a true amoeba does.

250 μm

(a) The brightly pigmented plasmodium, shown on a dead log, feeds on bacteria and other microorganisms.

FIGURE 26-19 The plasmodial slime mold Physarum polycephalum

(b) The reproductive structures are sporangia on stalks.

© Cabisco / Visuals Unlimited, Inc.

© Cabisco / Visuals Unlimited, Inc.

5

Fruiting body

50 μm

Spores

100 μm

6

Aggregation

Fruiting body 1

Slime mold amoebas

4

Slug differentiates

Aggregation

© Cabisco /Visuals Unlimited, Inc.

3

2

Multicellular slug forms

FIGURE 26-20 Animated The cellular slime mold Dictyostelium discoideum 50 μm

Slug

Slime mold amoebas ingest food, grow, and reproduce by cell division. After their food is depleted, hundreds of amoeboid cells stream together (inset in upper right) and form a migrating sluglike aggregate (inset in lower left). When it stops migrating, the aggregate forms a fruiting body on a stalk (inset in upper left). The fruiting body releases spores, each of which opens in a favorable environment to liberate an amoeboid cell.

When moisture or food becomes inadequate, certain cells send out a chemical signal, cyclic adenosine monophosphate (cAMP), that causes them to aggregate by the hundreds or thousands. During this stage the cells creep about for short distances as a single multicellular aggregate, or slug. Each cell of the slug retains its plasma membrane and individual identity. Eventually, the slug settles and reorganizes, forming a stalked fruiting body containing spores. After being released, each spore opens and a single haploid

amoeboid cell—the feeding stage—emerges. The spore-forming reproductive cycle is asexual, although sexual reproduction is observed occasionally. The life cycles of most cellular slime molds lack a flagellate stage. The cellular slime mold Dictyostelium discoideum is a model organism for the study of cell differentiation, cell communication, and cell motility and adhesion. Its biology has been studied intensively, particularly as it relates to cell signaling molecules, such

as cAMP, which are found in many organisms in addition to the cellular slime molds (see Chapter 6).

Flagellum Collar of microvilli

Choanoflagellates are closely related to animals Choanoflagellates are collared flagellates. These small, inconspicuous unikonts are found globally in both freshwater and marine environments. Choanoflagellates include both free-swimming and sessile species that are permanently attached by a thin stalk to bacteria-rich debris. Their single flagellum is surrounded at the base by a delicate collar of microvilli that trap food (FIG. 26-21). Choanoflagellates are of special interest because of their striking resemblance to collar cells in sponges (see Fig. 31-1b). Other animal phyla, such as cnidarians, flatworms, and echinoderms, also contain choanoflagellate-like cells, but no other group of protists has been observed to possess these cells. Evidence supporting the close relationship between choanoflagellates and animals includes comparative DNA sequence data of mitochondrial and nuclear genes. Given the similarities in structure and molecular genomics that has accumulated in recent years, many biologists hypothesize that choanoflagellates are the closest living non-animal relative of animals. Thus, living choanoflagellates and animals probably share a common choanoflagellate-like ancestor.

Review Why don’t we classify amoebas that have lobose pseudopodia with the rhizarians? Which group of unikonts is structurally and genetically most similar to animals?

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Cell

Lorica (protective cover)

Stalk

FIGURE 26-21 Choanoflagellates Choanoflagellates are free-living flagellates that obtain food by waving their flagella, causing water currents to carry bacteria and other small particles of food into the collar of microvilli. A colonial form is shown. Each cell is 5 to 10 mm long, not including the flagellum.

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

26.1 (page 538) 1 Discuss in general terms the diversity inherent in protists, including means of locomotion, modes of nutrition, interactions with other organisms, habitats, and modes of reproduction. Protists have various means of locomotion, including pseudopodia, ■ flagella, and cilia; a few are nonmotile.

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Protists obtain their nutrients autotrophically or heterotrophically. Protists are free-living or symbiotic, with symbiotic relationships ranging from mutualism to parasitism. Most protists live in the ocean or in freshwater ponds, lakes, and streams. Parasitic protists live in the body fluids or cells of their hosts. Many protists reproduce both sexually and asexually; others reproduce only asexually.

26.2 (page 539) 2 Discuss the hypothesis of serial endosymbiosis and briefly explain some of the evidence that supports it.

According to the hypothesis of serial endosymbiosis, mitochondria and chloroplasts arose from symbiotic relationships between larger cells and the smaller bacteria that were incorporated and lived within them. ■ Chloroplasts of red algae, green algae, and plants probably arose in a single primary endosymbiotic event in which a cyanobacterium was incorporated into a cell. Multiple secondary endosymbioses led to chloroplasts in euglenoids, dinoflagellates, diatoms, golden algae, and brown algae and to the nonfunctional chloroplasts in apicomplexans. 3 Describe the kinds of data biologists use to classify eukaryotes. ■ Relationships among protists are determined largely by ultrastructure, which is the fine details of cell structure revealed by electron microscopy, and by comparative molecular data. Biologists have compared nuclear genes, many of which code for proteins, in different protist taxa. ■

26.3 (page 540) 4 Summarize the basic features of excavates and distinguish among diplomonads, parabasilids, and euglenoids. ■ Excavates are a diverse group of unicellular protists with flagella, an excavated oral groove, and atypical, greatly modified mitochondria. The inclusion of diplomonads, parabasalids, and euglenoids in the excavate superfamily is controversial. ■ Diplomonads are excavates, organisms with a deep, or excavated, oral groove. Diplomonads have one or two nuclei, no functional mitochondria, no Golgi complex, and up to eight flagella.

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Parabasilids are anaerobic, flagellated excavates that often live in animals. Trichonymphs and trichomonads are examples of parabasilids. Euglenoids are unicellular and flagellate. Some euglenoids are photosynthetic.

26.4 (page 544)

■

26.5 (page 551) 8 Describe the forams and actinopods, and explain why many biologists

5 Contrast the two main groups of chromalveolates: alveolates and stramenopiles. ■ Chromalveolates probably originated as a result of secondary endosymbiosis in which an ancestral cell engulfed a red alga. Some DNA sequence data suggest the chromalveolates are not monophyletic. ■ Alveolates have similar ribosomal DNA sequences and alveoli, flattened vesicles located just inside the plasma membrane. Most stramenopiles have motile cells with two flagella, one of which has tiny hairlike projections off the shaft. 6 Distinguish among the alveolates: dinoflagellates, apicomplexans, and ciliates. ■ Dinoflagellates are mostly unicellular, biflagellate, photosynthetic alveolates of great ecological importance as producers in marine ecosystems. Their alveoli, flattened vesicles under the plasma membrane, often contain cellulose plates impregnated with silicates. Some dinoflagellates produce toxic blooms known as red tides. ■ Apicomplexans are parasites that produce sporozoites and are nonmotile. An apical complex of microtubules attaches the apicomplexan to its host cell. The apicomplexan Plasmodium causes malaria.

classify them in the rhizarian supergroup. ■ Rhizarians are amoeboid cells that often have hard outer shells, called tests, through which cytoplasmic projections extend; molecular evidence indicates this group is monophyletic. ■ Forams secrete many-chambered tests with pores through which cytoplasmic projections extend to move and obtain food. ■ Actinopods are mostly marine plankton that obtain food by means of axopods, slender cytoplasmic projections that extend through pores in their shells.

26.6 (page 552) 9 Describe evidence supporting the hypothesis that red algae and green algae should be included in a monophyletic group with land plants. ■ Red algae, green algae, and land plants, collectively called archaeplastids, are considered a monophyletic group based on molecular data and on the presence of chloroplasts bounded by outer and inner membranes. ■ Red algae, which are mostly multicellular seaweeds, are ecologically important in warm tropical ocean waters. ■ Green algae exhibit a wide diversity in size, structural complexity, and reproduction. Botanists hypothesize that ancestral green algae gave rise to land plants.

Watch the life cycle of the malaria parasite by clicking on the figure in CengageNOW. ■

Brown algae are multicellular stramenopiles that are ecologically important in cooler ocean waters. The largest brown algae (kelps) possess leaflike blades, stemlike stipes, anchoring holdfasts, and gas-filled bladders for buoyancy.

Ciliates are alveolates that move by hairlike cilia, have micronuclei (for sexual reproduction) and macronuclei (for controlling cell metabolism and growth), and undergo a sexual process called conjugation.

Watch the life cycles of a red alga and a green alga by clicking on the figures in CengageNOW.

26.7 (page 554) 10 Briefly describe and compare the following unikonts: amoebas, plasmodial slime molds, cellular slime molds, and choanoflagellates. Unikonts, which include fungi and animals, share a single posterior flagellum in flagellate cells. ■ Amoebas move and obtain food by phagocytosis, using cytoplasmic extensions called pseudopodia. ■ The feeding stage of plasmodial slime molds is a multinucleate plasmodium. Reproduction is by haploid spores produced within sporangia. ■ Cellular slime molds feed as individual amoeboid cells. They reproduce by aggregating into an aggregate (slug), then forming asexual spores. ■

Watch conjugation by clicking on the figure in CengageNOW.

7 Distinguish among the stramenopiles: water molds, diatoms, golden algae, and brown algae. Water molds have a coenocytic mycelium. They reproduce asexually by forming biflagellate zoospores and sexually by forming oospores. ■ Diatoms are mostly unicellular, with shells containing silica. Some diatoms are part of floating plankton, and others live on rocks and sediments where they move by gliding. Golden algae are mostly unicellular, biflagellate freshwater and ■ marine stramenopiles that are of ecological importance as a component of the ocean’s extremely minute nanoplankton. ■

Watch the life cycle of a cellular slime mold by clicking on the figures in CengageNOW. ■

Choanoflagellates are unikonts that are probably the closest living non-animal relative of animals. A collar of microvilli surrounds their single flagellum at the base.

T E S T YO U R U N D E R S TA N D I N G 1. Which of the following is not true of the protists? (a) they are unicellular, colonial, coenocytic, or simple multicellular organisms (b) their cilia and flagella have a 9 + 2 arrangement of microtubules (c) they are prokaryotic, as bacteria and archaea are (d) some are free-living, and some are endosymbionts (e) most are aquatic and live in the ocean or in freshwater ponds

2. Molecular evidence supports the view that all plastids evolved from an ancient (a) cyanobacterium (b) archaean (c) diplomonad (d) apicomplexan (e) unikont 3. Forams (a) are endosymbionts that live in many marine invertebrates (b) were responsible for the Irish potato famine in the 19th century (c) secrete many-chambered tests with pores through

which cytoplasmic extensions project (d) have numerous axopods that may aid in trapping and holding prey (e) have numerous cilia to direct food into the oral groove

12. Label the diagrams. Use Figures 26-5b and 26-8b to check your answers.

4. Paramecium and other ciliates often display a sexual phenomenon called (a) oogamy (b) conjugation (c) serial endosymbiosis (d) red tide (e) alternation of generations 5. Parasitic alveolates that form spores at some stage in their life belong to which group? (a) actinopods (b) ciliates (c) coccolithophorids (d) apicomplexans (e) dinoflagellates 6. Malaria (a) is transmitted by the bite of a female tsetse fly (b) is caused by a parasitic flagellate, Giardia intestinalis (c) is a serious form of amoebic dysentery caused by Entamoeba histolytica (d) is caused by an apicomplexan that spends part of its life cycle in the Anopheles mosquito and part in humans (e) is transmitted when people drink water tainted by a red tide 7. Photosynthetic protists with shells composed of two halves that fit together like a petri dish are (a) golden algae (b) diatoms (c) euglenoids (d) brown algae (e) forams 8. The pigments, energy reserve products, and cell walls found in land plants are also characteristic of (a) green algae (b) brown algae (c) golden algae (d) diatoms (e) euglenoids and diatoms 9. Kelps are _____ with multicellular bodies differentiated into blades, stipes, holdfasts, and gas-filled floats. (a) golden algae (b) diatoms (c) euglenoids (d) brown algae (e) red algae 10. Cellular slime molds (a) include Physarum and Phytophthora (b) are more closely related to bacteria than are any other protists (c) are responsible for late blight of potatoes, which led to starvation in Ireland in the 1840s (d) have double-gene fusion (e) form a slug when cells aggregate in response to cyclic AMP 11. Water molds reproduce asexually by forming _____ and sexually by forming _____. (a) oospores; holdfasts (b) zoospores; zooxanthellae (c) zoospores; oospores (d) holdfasts; oospores (e) oospores; zoospores

1. EVOLUTION LINK. Why are the protists considered paraphyletic? Use Figure 26-3 to help explain your answer. 2. ANALYZING DATA. Where on Figure 26-3 would you place the common ancestor of brown algae and water molds? The common ancestor of animals and fungi? 3. ANALYZING DATA. In the darkfield LM of unicellular choanoflagellates (at right), actin has been stained red, tubulin green, and DNA blue. Identify the cell parts that contain each of these chemical materials. 4. SCIENCE, TECHNOLOGY, AND SOCIETY. Why has resistance to traditional antimalarial drugs emerged in Plasmodium?

Nicole King Lab, University of California, Berkeley

CRITICAL THINKING

Additional questions are available in CengageNOW at www.cengage.com/ login.

27

Seedless Plants

Mark Edwards/Peter Arnold Inc.

Land plants. Note the ferns in the foreground and the moss-covered dead tree. Both mosses and ferns are seedless plants.

KEY CONCEPTS 27.1 Adaptations of land plants include a waxy cuticle to prevent water loss; multicellular gametangia; stomata; and for most plants, vascular tissues containing lignin. Plants undergo an alternation of generations between multicellular gametophyte and sporophyte generations. 27.2 Mosses and other bryophytes lack vascular tissues

A

bout 445 million years ago (mya), planet Earth would have seemed an inhospitable place because, although life abounded in the ocean,

it did not yet exist in abundance on land. Occasionally, perhaps, an animal would crawl out of the water onto land, but it never stayed there permanently because there was little to eat on land—not a single blade of grass, no fruit, and no seeds. During the next 30 million years, a time corresponding roughly to

and do not form true roots, stems, or leaves.

the Silurian period in geologic time (see Table 21-1), plants appeared in

27.3 In club mosses and ferns, lignin-hardened vascular

abundance and colonized the land. Where did they come from? Although

tissues that transport water and dissolved substances throughout the plant body have evolved.

plants living today exhibit great diversity in size, form, and habitat (see photograph), biologists hypothesize that they all evolved from a common ancestor that was an ancient green alga. Modern green algae share many biochemical and metabolic traits with modern plants. Both green algae and plants contain the same photosynthetic pigments: chlorophylls a and b and carotenoids, including xanthophylls (yellow pigments) and carotenes (orange pigments). Both store excess carbohydrates as starch and have cellulose as a major component of their cell walls. In addition, plants and some green algae share certain details of cell division, including formation of a cell plate during cytokinesis (see Chapter 10).

What exactly is a plant? A plant is a complex multicellular eukaryote that has cellulose cell walls, chlorophylls a and b in plastids,

released before fertilization, whereas in others the fertilized egg is released.

starch as a storage product, and may have cells with two anterior flagella. In addition, all plants develop from multicellular embryos that are enclosed in maternal tissues; this last character is one that distinguishes plants from green algae. Many botanists refer to land plants as embryophytes because of this character. Plants range in size from minute, almost microscopic duckweeds and water-meal to massive giant sequoias, some of the largest organisms that have ever lived. Plants include hundreds of thousands of species that live in varied habitats, from frozen arctic tundra to lush tropical rain forests to harsh deserts to moist stream banks.

27.1 ADAPTATIONS OF PLANTS TO LIFE ON LAND ■ ■ LEARNING OBJECTIVES 1 2

Discuss some environmental challenges of living on land and describe how several plant adaptations meet these challenges. Name the green algal group from which plants are hypothesized to have descended and describe supporting evidence.

The plant life cycle alternates between haploid and diploid generations Plants have a clearly defined alternation of generations in which they spend part of their lives in a multicellular haploid stage and part in a multicellular diploid stage (FIG. 27-2).1 The haploid portion of the life cycle is called the gametophyte generation because it gives rise to haploid gametes by mitosis. When two gametes fuse, the diploid portion of the life cycle, called the sporophyte generation, begins. The sporophyte generation produces haploid spores by the process of meiosis; these spores represent the first stage in the gametophyte generation. Let us examine alternation of generations more closely. The haploid gametophytes produce male gametangia, known as antheridia (sing., antheridium), in which sperm cells form, and/or female gametangia, known as archegonia (sing., archegonium), each bearing a single egg (FIG. 27-3). Sperm cells reach the female gametangium in a variety of ways, and one sperm cell fertilizes the egg to form a zygote, or fertilized egg. The diploid zygote is the first stage in the sporophyte generation. The zygote divides by mitosis and develops into a multicellular embryo, the young sporophyte plant. Embryo development takes place within the archegonium; thus, the embryo is protected as it develops. Eventually, the embryo grows into a mature sporophyte plant. The mature sporophyte has special cells called sporogenous cells (spore-producing cells, also called spore mother cells) that divide by meiosis to form haploid spores.

How were plants modified during the transition from life in the water to life on land? What are some features of plants that have let them colonize so many types of environments? One important difference between plants and algae is that a waxy cuticle covers fertilization of egg by sperm cell ¡ zygote ¡ embryo ¡ the aerial portion of a plant. Essential for existence on land, the mature sporophyte plant ¡ sporogenous cells ¡ meiosis cuticle helps prevent desiccation, or drying out, of plant tissues by ¡ spores evaporation. Plants that are adapted to moister habitats may have a very thin layer of wax, whereas those adapted to drier environments often have a thick, crusty cuticle. (Many desert plants also 1 have a reduced surface area, particularly of leaves, which minimizes For convenience, we limit our discussion to plants that are not polyploid, alwater loss.) though polyploidy is very common in land plants. We therefore use the terms Plants obtain the carbon they need for photosynthesis from diploid and 2n (and haploid and n) interchangeably, although these terms are the atmosphere as carbon dioxide (CO2). To be fixed into organic not actually synonymous. molecules such as sugar, CO2 must first diffuse into the chloroplasts that are inside green plant cells. Because a Developing waxy cuticle covers the external surfaces of leaves and gametes Sterile cells stems, however, gas exchange through the cuticle between the atmosphere and the interior of cells is negligible. Tiny pores called stomata (sing., stoma), which dot the surfaces of leaves and stems of almost all plants, facilitate gas exchange. The sex organs, or gametangia (sing., gametangium), Multicellular gametangium of most plants are multicellular, whereas the gametangia Unicellular of algae are unicellular (FIG. 27-1). Each plant gametangametangium gium has a layer of sterile (nonreproductive) cells that (a) (c) (b) (d) surrounds and protects the delicate gametes (eggs and sperm cells). In plants, the fertilized egg develops into a FIGURE 27-1 Generalized reproductive structures of algae and plants multicellular embryo (young plant) within the female (a, b) In algae, gametangia are generally unicellular. When the gametes are released, gametangium. Thus, the embryo is protected during its only the wall of the original cell remains. (c, d) In plants, the gametangia are multidevelopment. In algae, the fertilized egg develops away cellular, but only the inner cells become gametes. The gametes are surrounded by a from its gametangium; in some algae, the gametes are protective layer of sterile (nonreproductive) cells.

KEY POINT

Plants have an alternation of generations, spending part of the cycle in a haploid gametophyte stage and part in a diploid sporophyte stage. 1

The gametophyte produces gametes by mitosis. Gametophyte

5

Each spore has the potential to undergo mitosis and develop into a gametophyte.

Spore

Two haploid gametes fuse to form a zygote.

3

The zygote (fertilized egg) develops by mitosis into the sporophyte.

Egg

Sperm HAPLOID (n) GAMETOPHYTE GENERATION

Meiosis 4

2

Fertilization DIPLOID (2n) SPOROPHYTE GENERATION

Special cells of the sporophyte undergo meiosis to form spores.

Zygote Embryo

Sporophyte

FIGURE 27-2 The basic plant life cycle Depending on the plant group, the haploid or the diploid stage may be greatly enlarged or reduced; as you will learn, this fact has evolutionary significance.

All plants produce spores by meiosis, in contrast with algae and fungi, which may produce spores by meiosis or mitosis. The spores represent the first stage in the gametophyte generation. Each spore divides by mitosis to produce a multicellular gametophyte, and the cycle continues. Plants therefore alternate between a haploid gametophyte generation and a diploid sporophyte generation.

archegonia ¡ eggs mature ¡ spores ¡ gametophyte plants ¡ antheridia ¡ sperm cells

Developing sperm cells Sterile cells

Antheridium

Egg Sterile cells

(a) Each antheridium, the male gametangium, produces numerous sperm cells.

FIGURE 27-3 Plant gametangia Shown are generalized moss gametangia.

Archegonium

(b) Each archegonium, the female gametangium, produces a single egg.

Four major groups of plants exist today Structural and molecular data indicate that land plants probably descended from a group of green algae called charophytes or stoneworts (see Fig. 26-15d, which shows a charophyte). Recall from Chapter 26 that red algae, green algae, and land plants are collectively classified as archaeplastids. Molecular comparisons, particularly of DNA and RNA sequences, provide compelling evidence that green algae are closely allied to plants (FIG. 27-4). These comparisons among plants and various green algae include sequences of chloroplast DNA, certain nuclear genes, and ribosomal RNA. In each case, the closest match occurs between charophytes and plants, indicating that modern charophytes and plants probably share a recent common ancestor. Plants consist of four major groups: bryophytes, seedless vascular plants, and two groups of seeded vascular plants: gymnosperms and angiosperms (flowering plants) (FIG. 27-5; see also TABLE 27-1, which is an overview of the ten extant [living] plant phyla). The mosses and other bryophytes are small nonvascular plants that lack a specialized vascular, or conducting, system to trans-

port nutrients, water, and essential minerals (inorganic nutrients) throughout the plant body. In the absence of such a system, bryophytes rely on diffusion and osmosis to obtain needed materials. This reliance means that bryophytes are restricted in size; if they were much larger, some of their cells could not obtain enough necessary materials. Bryophytes do not form seeds, the reproductive structures discussed in Chapter 28. Bryophytes reproduce and disperse primarily via haploid spores. Recent molecular and fossil evidence, discussed later in the chapter, suggests that bryophytes may have been the earliest plants to colonize land. The other three groups of plants—seedless vascular plants, gymnosperms, and flowering plants—have vascular tissues and are thus known as vascular plants. The two vascular tissues are xylem, for conducting water and dissolved minerals, and phloem, for conducting dissolved organic molecules such as sugar. A key step in the evolution of vascular plants was the ability to produce lignin, a strengthening polymer in the walls of cells that function for support and conduction (see Chapter 33 for a discussion of plant cell wall chemistry, including lignin). The stiffening property of lignin enabled plants to grow tall, which let them maximize light

Archaeplastids

Green algae, including charophytes

Red algae

Land plants

Development of multicellular sporophyte embryo within maternal gametophyte tissues

Loss of red pigment, phycoerythrin

Ancestral eukaryote

Development of chloroplast from incorporated cyanobacterium

FIGURE 27-4 Evolution of land plants Cladograms such as this one represent an emerging consensus that is open to change as new discoveries are made.

KEY POINT

Based on recent evidence, scientists are beginning to reach a consensus about the evolutionary relationships among living plants. Land Plants VASCULAR SEED PLANTS

Angiosperms

Gymnosperms

Ferns

VASCULAR SEEDLESS PLANTS

Club mosses

Hornworts

Mosses

NONVASCULAR BRYOPHYTES

Liverworts

interception. The successful occupation of the land by plants, in turn, made the evolution of terrestrial animals possible by providing them with both habitat and food. Club mosses and ferns (which include whisk ferns and horsetails) are seedless vascular plants that, like the bryophytes, reproduce and disperse primarily via spores. Seedless vascular plants arose and diversified during the Silurian and Devonian periods of the Paleozoic era, between 444 mya and 359 mya. Club mosses and ferns extend back more than 420 million years and were of considerable importance as Earth’s dominant plants in past ages. Fossil evidence indicates that many species of these plants were the size of immense trees. Most living representatives of club mosses and ferns are small. The gymnosperms are vascular plants that reproduce by forming seeds. Gymnosperms produce seeds borne exposed (unprotected) on a stem or in a cone. Plants with seeds as their primary means of reproduction and dispersal first appeared about 359 mya, at the end of the Devonian period. These early seed plants diversified into many varied species of gymnosperms. The most recent plant group to appear is the flowering plants, or angiosperms, which arose during the early Cretaceous period of the Mesozoic era, about 130 mya. Like gymnosperms, flowering plants reproduce by forming seeds. Flowering plants, however, produce seeds enclosed within a fruit.

Evolution of seeds Evolution of dominant sporophyte, vascular tissue

Green algal ancestor

Evolution of cuticle, multicellular gametangia, multicellular embryos

FIGURE 27-5 Animated Plant evolution

Review ■

■

■

■

The four major groups of plants are bryophytes, seedless vascular plants, and two groups of seed plants—gymnosperms and angiosperms. The seed plants are covered in Chapter 28.

What are the most important environmental challenges that plants face living on land? What adaptations do plants have to meet these environmental challenges? From which group of green algae are plants hypothesized to have descended? Draw a simple diagram illustrating alternation of generations in plants.

27.2 BRYOPHYTES ■ ■ LEARNING OBJECTIVES 3 Summarize the features that distinguish bryophytes from other plants. 4 Name and briefly describe the three phyla of bryophytes. 5 Describe the life cycle of mosses and compare their gametophyte and sporophyte generations.

The bryophytes (from the Greek words meaning “moss plant”) consist of about 16,000 species of mosses, liverworts, and hornworts;

bryophytes are the only living nonvascular plants (TABLE 27-2). Because they have no means for extensive internal transport of water, sugar, and essential minerals, bryophytes are typically small. They generally require a moist environment for active growth and reproduction, but some bryophytes tolerate dry areas. The bryophytes are divided into three distinct phyla: mosses (phylum Bryophyta), liverworts (phylum Hepatophyta), and hornworts (phylum Anthocerophyta). These three groups differ in many ways and may or may not be closely related. They are usually studied together because they lack vascular tissues and have similar life cycles.

Moss gametophytes are differentiated into “leaves” and “stems” Mosses (phylum Bryophyta), which include about 9900 species, usually live in dense colonies or beds (FIG. 27-6a). Each individual

rate sexes: male plants that bear antheridia and female plants that bear archegonia. Other mosses produce anNonvascular plants with a dominant gametophyte generation theridia and archegonia on the same plant. (bryophytes) Fertilization occurs when one of the sperm cells fuses Phylum Hepatophyta (liverworts) with the egg within the archegonium. Sperm cells, which Phylum Bryophyta (mosses) have flagella, are transported from antheridium to archePhylum Anthocerophyta (hornworts) gonium by flowing water, such as splashing rain droplets. A raindrop lands on the top of a male gametophyte, and Vascular plants with a dominant sporophyte generation sperm cells are released into it from the antheridia. AnSeedless plants other raindrop landing on the male plant may splash the Phylum Lycopodiophyta (club mosses) sperm-laden droplet into the air and onto the top of a Phylum Pteridophyta (ferns and their allies, nearby female plant. Alternatively, arthropods such as inthe whisk ferns and horsetails) sects and mites may touch the sperm-laden fluid and inadvertently carry it for considerable distances. Once in a Seed plants film of water on the female moss, a sperm cell swims into Plants with naked seeds (gymnosperms) the archegonium, which secretes chemicals to attract and Phylum Coniferophyta (conifers) guide the sperm cells, and fuses with the egg. Phylum Cycadophyta (cycads) The diploid zygote, formed by fertilization, grows Phylum Ginkgophyta (ginkgoes) by mitosis into a multicellular embryo that develops into Phylum Gnetophyta (gnetophytes) a mature moss sporophyte. This sporophyte grows out Seeds enclosed within a fruit (angiosperms) of the top of the female gametophyte and remains atPhylum Anthophyta (angiosperms or flowering plants) tached and nutritionally dependent on the gametophyte Class Eudicotyledones (eudicots) throughout its existence (FIG. 27-6b). Initially green and Class Monocotyledones (monocots) photosynthetic, the sporophyte becomes golden brown at maturity. It consists of three main parts: a foot, which anchors the sporophyte to the gametophyte and absorbs minerals and nutrients from it; a seta, or stalk; and a capsule, which contains sporogenous cells (spore mother cells). The plant has tiny, hairlike absorptive structures called rhizoids and an capsule of some species is covered by a caplike structure, the calypupright, stemlike structure that bears leaflike blades, each normally tra, which is derived from the archegonium. consisting of a single layer of undifferentiated cells except at the The sporogenous cells undergo meiosis to form haploid spores midrib. Because mosses lack vascular tissues, they do not have true (see Fig. 27-7). When the spores are mature, the capsule opens and roots, stems, or leaves; the moss structures are not homologous releases the spores, which are then transported by wind or rain. If a to roots, stems, or leaves in vascular plants. Some moss species moss spore lands in a suitable spot, it germinates and grows into a have water-conducting cells and sugar-conducting cells, although filament of cells called a protonema. The protonema, which superfithese cells are not lignified or as specialized or effective as the concially resembles a filamentous green alga, forms buds, each of which ducting cells of vascular plants. grows into a green gametophyte, and the life cycle continues. Alternation of generations is clearly defined in the life cycle Biologists consider the haploid gametophyte generation the of mosses (FIG. 27-7). The green moss gametophyte often bears its gametangia at the top of the plant. Many moss species have sepadominant generation in mosses because it lives independently TABLE 27-1

Ten Extant Plant Phyla

TABLE 27-2

A Comparison of Major Groups of Seedless Plants

Plant Group

Dominant Stage of Life Cycle

Nonvascular; reproduce by spores (bryophytes) Liverworts (phylum Hepatophyta) Gametophyte: thalloid or leafy plant Mosses (phylum Bryophyta) Gametophyte: leafy plant Hornworts (phylum Anthocerophyta) Vascular; reproduce by spores Club mosses (phylum Lycopodiophyta) Ferns (phylum Pteridophyta) Whisk ferns (phylum Pteridophyta) Horsetails (phylum Pteridophyta)

Gametophyte: thalloid plant

Sporophyte: roots, rhizomes, erect stems, and leaves (microphylls) Sporophyte: roots, rhizomes, and leaves (megaphylls) Sporophyte: rhizomes and erect stems; no true roots or leaves Sporophyte: roots, rhizomes, erect stems, and leaves (reduced megaphylls)

Representative Genera

Marchantia Polytrichum, Sphagnum, Physcomitrella Anthoceros

Lycopodium, Selaginella Pteridium, Polystichum, Azolla, Platycerium Psilotum Equisetum

Rod Planck/Dembinsky Photo Associates

Vascular seed plants

Vascular seedless plants

Nonvascular bryophytes

Green algal ancestor

thin layer of organic matter in which grasses and other plants can grow. Because they grow tightly packed in dense colonies, mosses hold soil in place and help prevent erosion. At the same time, they retain moisture that they and other organisms need. Waxwings and other birds use moss, along with twigs and grass, as nesting material. Commercially, the most important mosses are the peat mosses in the genus Sphagnum. One of the distinctive features of Sphagnum “leaves” is the presence of many large, empty (a) Moss gametophytes. Close-up of haircap cells that absorb and hold water. moss (Polytrichum commune) gametophytes. This feature makes peat moss a useThe haircap moss is a popular ground cover in rock gardens, particularly in Japan. ful packing material for shipping live plants as well as a good soil conditioner. Added to sandy soils, for example, peat moss helps absorb and retain moisture. The acidic and anaerobic conditions of a peat bog retard the Capsule growth of bacterial and fungal decomposers. As a result, dead peat mosses accumulate as thick deposits—some several meters in depth—under the growing mat of living peat mosses. Over time, the organic material compresses to form peat. In some countries, Seta such as Ireland and Scotland, people cut out blocks of peat that has accumulated for hundreds of years in peat bogs, dry them, and burn them for fuel. Occasionally, the remains of well-preserved humans have been uncovered during the excavations of old peat bogs in Ireland and other parts of Europe (FIG. 27-8). The name moss is often misused to refer to plants that are not truly mosses. For example, reindeer “moss” is a lichen that is a dominant form of vegetation in the arctic tundra, Spanish “moss” Foot is a flowering plant, and club “moss” (discussed later in the chapter) is a relative of ferns.

David Cavagnaro

Liverwort gametophytes are either thalloid or leafy

(b) Moss sporophytes. Each consisting of a foot, seta, and capsule, the haircap moss sporophytes grow out of the top of the gametophytes. Spores are produced within the capsule.

FIGURE 27-6 Bryophytes: mosses

of the diploid sporophyte. In contrast, the moss sporophyte is attached to and nutritionally dependent on the gametophyte. Mosses make up an inconspicuous but significant part of their environment. They play an important role in forming soil. Mosses, which form mats that cover the rock, eventually die, forming a

Liverworts (phylum Hepatophyta) consist of about 6000 species of nonvascular plants with a dominant gametophyte generation, but the gametophytes of some liverworts are quite different from those of mosses. Their body form is often a flattened, lobed structure called a thallus (pl., thalli) that is not differentiated into leaves, stems, or roots. The common liverwort, Marchantia polymorpha, is thalloid (FIGS. 27-9a and b). Liverworts are so named because the lobes of their thalli superficially resemble the lobes of the human liver; wort is derived from the Old English word wyrt, meaning “plant.” On the underside of the liverwort thallus are hairlike rhizoids that anchor the plant to the soil. Other liverworts, known as leafy liverworts, superficially resemble mosses, with leaflike blades, “stems,” and rhizoids rather than a lobed thallus (FIG. 27-9c). As in the mosses, leafy liverwort “leaves” consist of a single layer of undifferentiated cells. Like other bryophytes, liverworts are small, generally inconspicuous plants that are largely restricted to

KEY POINT

The dominant phase in the life cycle of mosses, like that of other bryophytes, is the gametophyte.

1 Antheridia and/or archegonia

form at the tip of gametophyte shoots.

Gametophyte plants Buds on protonema

Antheridia with sperm cells

6 When a haploid spore

germinates, it forms a protonema.

Spores released

Protonema Sperm cell Archegonium with egg

HAPLOID (n) GAMETOPHYTE GENERATION

Meiosis

2 Splashing raindrops

transfer sperm cells from antheridia to archegonia.

Fertilization

DIPLOID (2n) SPOROPHYTE GENERATION 4 The mature sporophyte is

Calyptra 5 Sporogenous cells

an unbranched stalk with a sporangium at its tip.

Zygote

Capsule

within each sporangium undergo meiosis to form spores.

3 The fertilized egg grows

Sporophyte

Gametophyte plant

by mitosis to form the embryonic sporophyte.

Embryo

FIGURE 27-7 Animated The life cycle of mosses The gametophyte generation is dominant in the moss life cycle. After sexual reproduction, the sporophyte grows out of the gametophyte.

▲

© Robin Weaver/Alamy

damp environments. Unlike mosses, hornworts, and other plants, liverworts lack stomata, although some liverworts have surface pores thought to be analogous to stomata. Liverworts reproduce both sexually and asexually (see Figs. 27-9a and 27-9b). Their sexual reproduction involves production of archegonia and antheridia on the haploid gametophyte. In some liverworts, these gametangia are borne on stalked structures called archegoniophores, which bear archegonia, and antheridiophores, which bear antheridia. Their life cycle is basically the same as that of mosses, FIGURE 27-8 Preserved human remains in a peat bog in Denmark The clothing and features of the Tollund man, estimated to be about 2000 years old, are remarkably well preserved because the bog’s acidic conditions inhibited decay.

Antheridiophore © blickwinkel/Alamy

Gametophyte thallus

Antheridia with sperm cells

Gemmae cup

(a) Flattened, ribbon-like lobes characterize the gametophyte of the common liverwort (Marchantia polymorpha). This male gametophyte thallus has both asexual gemmae cups and sexual antheridiophores, which produce sperm-bearing antheridia.

Ed Reschke/Peter Arnold, Inc.

Archegoniophore

Gametophyte thallus

Achegonia with eggs

Following fertilization, a sporophyte develops, hanging upside-down on the archegoniophore.

(b) Achegoniophores on the female gametophyte thallus of M. polymorpha produce egg-bearing archegonia. Liverwort sporophytes, which are always attached to and dependent on the gametophyte plant, consists of a foot, seta, and capsule. Meiosis occurs in the capsule and produces haploid spores.

Foot

© Custom Medical Stock Photo/Alamy

Seta Tissue derived from archegonium

Capsule

Sporogenous cells that undergo meiosis

(c) Porella is a leafy liverwort. The leafy plant is the gametophyte. It bears antheridia and archegonia on special branches that look quite similar to the nonreproductive branches. After fertilization, a small sporophyte develops that produces spores following meiosis.

FIGURE 27-9 Bryophytes: liverworts

Sporophyte

although some of the structures look quite different. Splashing raindrops transport sperm cells to the archegonia, where fertilization takes place. The resulting zygote develops into a multicellular embryo that becomes a mature sporophyte. The liverwort sporophyte is attached to the gametophyte, as in mosses. Sporogenous cells in the capsule of the sporophyte undergo meiosis, producing haploid spores. Each spore has the potential to develop into a green gametophyte, and the cycle continues. Some liverworts reproduce asexually by forming tiny balls of tissue called gemmae (sing., gemma), which are borne in a saucershaped structure, the gemmae cup, directly on the liverwort thallus (see Fig. 27-9a). Splashing raindrops and small animals help disperse gemmae. When a gemma lands in a suitable place, it grows into a new liverwort thallus. Liverworts may also reproduce asexually by thallus branching and growth. The individual thallus lobes elongate, and each becomes a separate plant when the older part of the thallus that originally connected the individual lobes dies.

Mature sporangium splits open

Spores

Sporophyte

Hornwort gametophytes are inconspicuous thalloid plants Hornworts (phylum Anthocerophyta) are a small group of about 100 species of bryophytes whose gametophytes superficially resemble those of the thalloid liverworts. Hornworts live in disturbed habitats such as fallow fields and roadsides. Hornworts may or may not be closely related to other bryophytes. For example, their cell structure, particularly the presence of a single large chloroplast in each cell, resembles certain algal cells more than plant cells. In contrast, mosses, liverworts, and other plants have many disc-shaped chloroplasts per cell. In the common hornwort (Anthoceros natans), archegonia and antheridia are embedded in the gametophyte thallus rather than on archegoniophores and antheridiophores. After fertilization and development, the needlelike sporophyte projects out of the gametophyte thallus, forming a spike or “horn”—hence the name hornwort. A single gametophyte often produces multiple sporophytes (FIG. 27-10). Meiosis occurs, forming spores within each sporangium (pl., sporangia), or spore case. The sporangium splits open from the top to release the spores; each spore can give rise to a new gametophyte thallus. A unique feature of hornworts is that the sporophytes, unlike those of mosses and liverworts, continue to grow from their bases for the remainder of the gametophyte’s life, a characteristic known as indeterminate growth. Some botanists think that indeterminate growth may indicate that hornworts evolved from plants with larger, more complex sporophytes.

Bryophytes are used for experimental studies Botanists use certain bryophytes as experimental models to study many fundamental aspects of plant biology, including genetics, growth and development, plant ecology, plant hormones, and photoperiodism, which is plant responses to varying periods of night and day length. The moss Physcomitrella patens is a particularly important research organism for studying plant evolution because its features and genome can be compared to those of algae and flowering

Gametophyte with embedded archegonia and antheridia

FIGURE 27-10 Bryophytes: hornworts The gametophyte of the common hornwort (Anthoceros natans) is a small thallus with unicellular rhizoids on the lower (ventral) surface. After fertilization, sporophytes project up out of the gametophyte, forming “horns.”

plants (FIG. 27-11). In this regard, Physcomitrella is a plant equivalent of the fruit fly Drosophila, which is an important model organism for studies of animal inheritance, development, and evolution. As experimental organisms, Physcomitrella and other bryophytes are easy to grow on artificial media and do not require much space because they are so small.

Recap: details of bryophyte evolution are based on fossils and on structural and molecular evidence Plants are a monophyletic group—that is, all plants probably evolved from a common ancestral green alga. Fossil evidence indicates that bryophytes are ancient plants, probably the first group of plants to arise from the common plant ancestor. The fossil record of ancient bryophytes is incomplete, consisting mostly of spores and small tissue fragments, and can be interpreted in different ways. As a result, it does not provide a definite answer on bryophyte evolution. The oldest known recognizable plant fossils are dated at about 425 million years old. These fossils resemble modern liverworts in many respects, but the spores are virtually identical to those

KEY EXPERIMENT QUESTION: Does the Physcomitrella genome reveal any insights in plant evolution? HYPOTHESIS: Comparing the genome of the model organism Physcomitrella with genomes of other land plants as well as green algae will provide details of genes that were probably associated with the colonization of land. EXPERIMENT: Several dozen scientists in Germany, the United States, Japan, the United Kingdom, Australia, Canada, and Belgium collaborated to produce the draft genome sequence for the model organism Physcomitrella (see photograph). This bryophyte was selected because researchers have used Physcomitrella in a wide variety of experiments and have a good understanding of its biology.

in 470-million-year-old rocks. In 2003, scientists reported fossil fragments of tiny liverwort-like plants associated with ancient spores that were discovered in Oman. This evidence suggests that liverwort-like plants may have been the earliest plants to colonize land.

Review ■

■

Which of the following are parts of the gametophyte generation in mosses: antheridia, zygote, embryo, capsule, archegonia, sperm cells, egg cell, spores, and protonema? How are mosses, liverworts, and hornworts similar? How is each group distinctive?

27.3 SEEDLESS VASCULAR PLANTS ■ ■ LEARNING OBJECTIVES Courtesy of David J. Cove, University of Leeds

6 Discuss the features that distinguish seedless vascular plants from algae

After sequencing the genome of Physcomitrella, it was compared with available genomes of green algae (Ostreococcus and Chlamydomonas) and land plants—the model flowering plant Arabidopsis, rice (Oryza sativa), and western balsam poplar (Populus trichocarpa).

RESULTS AND CONCLUSION: Researchers found that Physcomitrella has lost some genes needed for life in the water and gained new genes needed for life on land. Every land plant has to meet the same environmental challenges, such as obtaining enough water and preventing excessive water loss, tolerating high and low temperatures, and adapting to increased levels of solar radiation. These challenges have resulted in many modifications in plant body plans and physiological processes. Bryophytes and vascular plants living today represent remnants of the first plants that colonized land some 450 mya. Compared to green algae, Physcomitrella lacks the genes needed to survive in aquatic environments but has gained genes involving plant signaling molecules (auxin, abscisic acid, and cytokinins) needed to adapt to drought, extremes of temperature, and light reception. Compared to Physcomitrella, other land plants have gained additional signaling molecules (gibberellins, jasmonic acid, and brassinosteroids) that enable a more complex tolerance of environmental stressors. Because of its intermediary position in evolution (i.e., between green algae and vascular land plants), Physcomitrella’s genome allows biologists to reconstruct which genes may have been acquired and lost during the colonization of land by the last common ancestor of all land plants. Source: Rensing, S. A., et al. “The Physcomitrella Genome Reveals Evolutionary Insights into the Conquest of Land by Plants.” Science, Vol. 319, Jan. 4, 2008.

FIGURE 27-11 The moss Physcomitrella, the first bryophyte to have its genome sequenced

7 8

and bryophytes. Name and briefly describe the two phyla of seedless vascular plants. Describe the life cycle of ferns and compare their sporophyte and gametophyte generations.

9 Compare the generalized life cycles of homosporous and heterosporous plants.

The most important adaptation found in seedless vascular plants, though absent in algae and bryophytes, is specialized vascular tissues—xylem and phloem—for support and conduction. This system of conduction lets vascular plants grow larger than bryophytes because water, minerals, and sugar are transported to all parts of the plant. Although seedless vascular plants in temperate environments are relatively small, tree ferns in the tropics may grow to heights of 18 m (60 ft). All seedless vascular plants have true stems with vascular tissues, and most also have true roots and leaves. Botanists have extensively studied the evolution of the leaf as the main organ of photosynthesis. The two basic types of true leaves—microphylls and megaphylls—evolved independently of each other (FIG. 27-12). The microphyll, which is usually small and has a single vascular strand, probably evolved from small, projecting extensions of stem tissue (enations). Only one group of living plants, the club mosses, has microphylls. In contrast, megaphylls probably evolved from stem branches that gradually filled in with additional tissue (webbing) to form most leaves as we know them today. Megaphylls have more than one vascular strand, as we would expect if they evolved from branch systems. Ferns (with the exception of whisk ferns, discussed later in the chapter), gymnosperms, and flowering plants have megaphylls. Recent evidence suggests megaphylls evolved over a 40million-year period in the Late Paleozoic era in response to a gradual decline in the level of atmospheric CO2. As CO2 declined, plants developed a flattened blade with more stomata for gas exchange. (More stomata allowed cells inside the leaf to get enough CO2.) There are two main clades of seedless vascular plants: the club mosses and the ferns. Biologists originally considered horsetails

Stem Microphyll Vascular tissue

Enation Vein

Smooth stem

Enation

Vascular supply to enation

Microphyll (one vein)

(a) Microphyll evolution. Microphylls probably originated as outgrowths (enations) of stem tissue that developed a single vascular strand later. Club mosses have microphylls.

Equal branches Vascular tissue

Dichotomously branching stem

Thicker main Dichotomous stem end branches

Thinner side branch

Overtopping (unequal branching)

Planation (branching in same plane)

Webbing of side branch system

Megaphyll (many veins)

(b) Megaphyll evolution. Megaphylls probably evolved from the evolutionary modification of side branches. Webbing is the evolutionary process in which the spaces between close branches become filled with chlorophyll-containing cells. Ferns, horsetails, gymnosperms, and flowering plants have megaphylls.

FIGURE 27-12 Evolution of microphylls and megaphylls

and whisk ferns distinct enough to be classified in separate phyla. However, many kinds of evidence, such as DNA comparisons and similarities in sperm structure, have resulted in their being reclassified as ferns. As shown in FIGURE 27-13, ferns, including horsetails and whisk ferns, are a monophyletic group and the closest living relatives of seed plants (also see Table 27-2).

Club mosses are small plants with rhizomes and short, erect branches Club mosses (phylum Lycopodiophyta) were important plants millions of years ago, when species that are now extinct often reached great size (FIG. 27-14a). These large, treelike plants were major contributors to our present-day coal deposits (see Inquiring About: Ancient Plants and Coal Formation). The 1200 or so species of club mosses living today, such as Lycopodium (FIG. 27-14b), are small (less than 25 cm, or 10 in, tall), attractive plants common in temperate woodlands. They possess true roots; both rhizomes and erect aerial stems; and small, scalelike leaves (microphylls). Sporangia are borne on reproductive leaves that are either clustered in conelike strobili at the tips

of stems or scattered in reproductive areas along the stem. Club mosses are evergreen and often fashioned into Christmas wreaths and other decorations. In some areas they are endangered by overharvesting. That common names are sometimes misleading in biology is vividly evident in this group of plants. The most common names for the phylum Lycopodiophyta are “club mosses” and “ground pines,” yet these plants are neither mosses, which are nonvascular, nor pines, which are seed plants.

Ferns are a diverse group of spore-forming vascular plants Most of the 11,000 species of ferns (phylum Pteridophyta) are terrestrial, although a few have adapted to aquatic habitats. Ferns range from the tropics to the Arctic Circle, with most species living in tropical rain forests, where they perch high in the branches of trees. In temperate regions, ferns commonly inhabit swamps, marshes, stream banks, and moist woodlands (FIG. 27-15a). Some species grow in fields, rocky crevices on cliffs or mountains, or even deserts.

KEY POINT

Seedless vascular plants include lycophytes and ferns.

SEEDLESS VASCULAR PLANTS

Seed plants

Horsetails

Whisk ferns

Ferns

FERNS

Club mosses and spike mosses

LYCOPHYTES

Loss of roots Vascular seed plants

Vascular seedless plants

Nonvascular bryophytes

The life cycle of ferns involves a clearly defined alternation of generations. The ferns grown as houseplants (such as Boston fern, maiden-hair fern, and staghorn fern) represent the larger, more conspicuous sporophyte generation. The fern sporophyte consists of a horizontal underground stem, or rhizome, that bears leaves, called fronds, and true roots. As each young frond first emerges from the ground, it is tightly coiled and resembles the top of a violin, hence the name fiddlehead (FIG. 27-15b). As fiddleheads grow, they unroll and expand to form fronds. Fern fronds are usually compound (the blade is divided into leaflets), with the leaflets forming beautifully complex leaves. Fronds, roots, and rhizomes all contain vascular tissues. Spore production usually occurs in certain areas on the fronds, which develop sporangia. Many species bear the sporangia in clusters, called sori (sing., sorus) (FIGS. 27-15c and d). Within sporan-

Evolution of seeds

Evolution of microphylls Evolution of megaphylls

Ancient vascular plant

Green algal ancestor

FIGURE 27-13 Evolutionary relationships among extant seedless vascular plants These relationships are based on structural and molecular comparisons.

Ed Reschke

Leaves (microphylls)

(a) Reconstruction of Lepidodendron, an ancient club moss the size of a large tree. Numerous fossils of Lepidodendron were preserved in coal deposits, particularly in Great Britain and the central United States. (Redrawn from Hirmer, M., Handbuch der Paläobotanik, R. Olderbourg,

(b) The sporophyte of Lycopodium, a club moss, has small, scalelike, evergreen leaves (microphylls). Spores are produced in sporangia on reproductive leaves clustered in a conelike strobilus (shown) or, in other species, scattered along the stem.

▲

Strobilus

gia, sporogenous cells (spore mother cells) undergo meiosis to form haploid spores. The sporangia burst open and discharge spores that may germinate and grow by mitosis into gametophytes. The mature fern gametophyte, which bears no resemblance to the sporophyte, is a tiny (less than half the size of one of your fingernails), green, often heart-shaped structure that grows flat against the ground. Called a prothallus (pl., prothalli), the fern gametophyte lacks vascular tissues and has tiny, hairlike absorptive rhizoids to anchor it (FIG. 27-15e). The prothallus usually produces both archegonia and antheridia on its underside. Each archegonium contains FIGURE 27-14 Seedless vascular plants: club mosses

In qu i ri n g A b o u t

How did the coal that industrial society depends on for energy form? Coal, which is burned to produce electricity and to manufacture items of steel and iron, is one of the most important fossil fuels. Although mined, coal is not an inorganic mineral like gold or aluminum but an organic material formed from the remains of ancient vascular plants, particularly those of the Carboniferous period, approximately 320 mya. Four main groups of plants contributed to coal formation. Two were seedless vascular plants: the club mosses and ferns (see figure), including horsetails. The other two were seed plants: seed ferns (now extinct) and early gymnosperms. It is hard to imagine that relatives of the small, relatively inconspicuous club mosses, ferns, and horsetails of today were so significant in forming vast beds of coal. However, many members of these groups that existed during the Carboniferous period were giants compared with their modern counterparts and formed immense forests. (Figure 21-12 shows a reconstruction of a Carboniferous forest.) The climate during the Carboniferous period was warm, moist, and mild. Plants

in most locations could grow year-round because of the favorable conditions. Forests of these plants often grew in low-lying, swampy areas that periodically flooded when the sea level rose. As the sea level receded, these plants would re-establish. When these large plants died or were blown over in storms, they decomposed incompletely because they were covered by swamp water. (The anaerobic conditions of the water prevented wood-rotting fungi from decomposing the plants, and anaerobic bacteria do not decompose wood rapidly.) Thus, over time the partially decomposed plant material accumulated and consolidated. Layers of sediment formed over the plant material each time the water level rose and flooded the low-lying swamps. With time, heat and pressure built up in these accumulated layers and converted the plant material to coal and the sediment layers to sedimentary rock. Much later, geologic upheavals raised the layers of coal and sedimentary rock. Coal is usually found in seams, underground layers that vary in thickness from 2.5 cm (1 in) to more than 30 m (100 ft).

a single egg, whereas numerous sperm cells are produced in each antheridium. Ferns use water as a transport medium. The flagellate sperm cells swim, usually from a nearby prothallus, to the neck of an archegonium through a thin film of water on the ground underneath the prothallus. After one of the sperm cells fertilizes the egg, a diploid zygote grows by mitosis into a multicellular embryo (an immature sporophyte). At this stage, the sporophyte embryo is attached to and dependent on the gametophyte; but as the embryo matures, the prothallus withers and dies, and the sporophyte becomes free-living. The fern life cycle alternates between the dominant, diploid sporophyte with its rhizome, roots, and fronds, and the haploid gametophyte (prothallus) (FIG. 27-16). The sporophyte generation is dominant not only because it is larger than the gametophyte but also because it persists for an extended period (most fern sporophytes are perennials), whereas the gametophyte dies soon after reproducing. Whisk ferns are classified as reduced ferns Only about 12 species of whisk ferns (phylum Pteridophyta) exist today, and the fossil record contains several extinct species. Whisk ferns, which live mainly in the tropics and subtropics, are relatively simple in structure and lack true roots and leaves but have vascularized stems. Psilotum nudum, a representative whisk fern, has both a horizontal underground rhizome and vertical

© David Lyons/Alamy

A N C I E N T P L A N T S A N D COA L F O R M AT I O N

This piece of Carboniferous coal contains a fossilized fern. The coal and fossil are about 300 million years old.

The various grades of coal (lignite, the lowest grade; sub-bituminous; bituminous; and anthracite, the highest grade) formed as a result of the different temperatures and pressures to which the layers were exposed. Coal exposed to high heat and pressure during its formation is drier, is more compact (and therefore harder), and has a higher heating value (that is, a higher energy content).

aerial stems (FIG. 27-17a).Whenever the stem forks, or branches, it always divides into two equal halves. Botanists call this forking dichotomous branching. In contrast, when most plant stems branch, one stem is more vigorous and becomes the main trunk. The upright stems of Psilotum are green and are the main organs of photosynthesis. Tiny, round sporangia, borne directly on the erect, aerial stems, contain sporogenous cells that undergo meiosis to form haploid spores. After dispersal, the spores germinate to form haploid prothalli. Because they grow underground, the prothalli of whisk ferns are difficult to study (FIG. 27-17b). They are nonphotosynthetic as a result of their subterranean location, and they apparently have a symbiotic relationship with mycorrhizal fungi that provides them with sugar and essential minerals (see Chapter 29). Botanists have carefully studied whisk ferns in recent years. Molecular data, including comparisons of nucleotide sequences of ribosomal RNA, chloroplast DNA, and mitochondrial DNA in living species, support the hypothesis that the whisk ferns should be classified as reduced ferns rather than as surviving representatives of extinct vascular plants (see discussion of rhyniophytes later in the chapter). Horsetails are an evolutionary line of ferns About 300 mya the horsetails (phylum Pteridophyta) were among the dominant plants and grew as large as modern trees (FIG. 27-18a).

John Arnaldi

© Cengage

Ed Reschke

(a) Fronds. The Christmas fern (Polystichum acrostichoides), photographed in the Great Smoky Mountains in Tennessee, has fronds that grow to 0.6 m (2 ft) in length.

(b) Fiddleheads. Some fiddleheads are edible.

(c) Sori. These round sori of rabbits foot fern (Polypodium aureum) are arranged in two prominent rows on the leaf’s underside.

© Carolina Biological Supply Company/Phototake. All rights reserved.

Prothallus

Ted Kinsman/Photo Researchers, Inc.

Archegonium

Sporangium Antheridium Rhizoid

(d) Sporangia. SEM of sporangia in a fern (Dryopteris filix-mas) sorus.

100 μm

(e) Prothallus. The prothallus is the gametophyte generation of a fern.

3 mm

FIGURE 27-15 Seedless vascular plants: ferns

Because they contributed to Earth’s vast coal deposits, these ancient horsetails, like ancient club mosses, are still significant today. The few surviving horsetails, about 15 species in the genus Equisetum, grow mostly in wet, marshy habitats and are less than 1.3 m (4 ft) tall, but extremely distinctive (FIG. 27-18b). They are widely distributed on every continent except Australia. Traditionally classified in their own phylum, horsetails are now grouped with ferns. This reclassification is based on molecular similarities between horsetails and other ferns. Horsetails have true roots, stems (both rhizomes and erect aerial stems), and small leaves. The hollow, jointed stems are impregnated with silica, which gives them a gritty texture. Small leaves, interpreted as reduced megaphylls, are fused in whorls at each node (the area on the stem where leaves attach). The green stem is the main organ of photosynthesis. Horsetails are so named

because certain vegetative (nonreproductive) stems have whorls of branches that give the appearance of a bushy horse’s tail. In the past, horsetails were called “scouring rushes” and were used to scrub pots and pans along stream banks. Each reproductive branch of a horsetail bears a terminal conelike strobilus (pl., strobili). The strobilus consists of several stalked, umbrella-like structures, each of which bears 5 to 10 sporangia in a circle around a common axis. The horsetail life cycle is similar in many respects to the fern life cycle. In horsetails, as in ferns, the sporophyte is the conspicuous plant, whereas the gametophyte is a minute, lobed thallus ranging in width from the size of a pinhead to about 1 cm across. The sporophyte and gametophyte are both photosynthetic and nutritionally independent at maturity. Like ferns, horsetails require water as a medium for flagellate sperm cells to swim to the egg.

KEY POINT

The dominant phase in the life cycle of ferns, like that of all other vascular plants, is the sporophyte.

Underside of enlarged mature gametophyte (prothallus) 4 Each spore has

the potential to germinate and develop into a prothallus, or fern gametophyte.

5 Gametes form

by mitosis. Egg

Archegonium Spores released

Rhizoids Antheridium 3 Sporogenous

cells within each sporangium undergo meiosis, forming spores.

Sperm cell

HAPLOID (n) GAMETOPHYTE GENERATION

Meiosis

Fertilization Zygote

DIPLOID (2n) SPOROPHYTE GENERATION

2 A sorus

contains a cluster of sporangia.

Frond Leaf of young sporophyte Leaf cross section 1 The fern

sporophyte develops reproductive structures on its leaves.

Haploid prothallus

Fiddlehead

Roots Underside of a frond

Rhizome

6 The sperm swims

to the archegonium and fertilizes the egg within, resulting in the start of the fern sporophyte generation.

Root of young sporophyte

Fern (mature sporophyte)

FIGURE 27-16 Animated The life cycle of ferns Note the clearly defined alternation of generations between the gametophyte (prothallus) and sporophyte (leafy plant) generations.

Some ferns and club mosses are heterosporous In the life cycles examined thus far, plants produce only one type of spore as a result of meiosis. This condition, known as homospory, is characteristic of bryophytes, horsetails, whisk ferns, and most ferns and club mosses. However, certain ferns and club mosses (known as spike mosses) exhibit heterospory, in which they produce two types of spores: microspores and megaspores. FIGURE 27-19 illustrates the generalized life cycle of a heterosporous plant. The sporophyte plant produces sporangia within a conelike strobilus. Each strobilus usually bears two kinds of

sporangia: microsporangia and megasporangia. Microsporangia are sporangia that produce microsporocytes (also called microspore mother cells), which undergo meiosis to form microscopic, haploid microspores. Each microspore develops into a male gametophyte that produces sperm cells within antheridia. Megasporangia produce megasporocytes (also called megaspore mother cells). When megasporocytes undergo meiosis, they form haploid megaspores, each of which develops into a female gametophyte that produces eggs in archegonia. The development of male gametophytes from microspores and of female gametophytes from megaspores occurs within their

▲

FIGURE 27-17 Whisk ferns

Antheridia

Sporangia

Archegonia Aerial stem with scalelike outgrowths (no leaves)

Rhizoids

John Arnaldi

(b) The gametophyte of the whisk fern Psilotum nudum. The gametophyte (prothallus) lives underground, nourished by mycorrhizal fungi.

(a) The sporophyte of the whisk fern Psilotum nudum. The stem is the main organ of photosynthesis in this rootless, leafless, vascular plant. Sporangia, which turn yellow as they mature, are borne on short lateral branches directly on the stems.

Strobilus

J. Robert Waaland/Biological Photo Service

Vegetative shoots

(b) Equisetum telematia, a horsetail with a wide distribution in Eurasia, Africa, and North America, has unbranched reproductive shoots bearing conelike strobili and separate, highly branched vegetative (nonreproductive) shoots. In some horsetail species, both reproductive and vegetative shoots are unbranched.

▲

(a) Reconstruction of Calamites, an ancient horsetail that grew as tall as many modern trees—to 20 m (about 65 ft). Calamites had an underground rhizome where roots and aerial shoots originated. (Redrawn from Emberger, L., Les Plantes Fossiles, Masson et Cie, Paris, 1968.)

Reproductive shoots

FIGURE 27-18 Horsetails

KEY POINT

Heterospory—the production of two types of spores—was the forerunner of the evolution of seeds.

2 Each megaspore may

develop into a female gametophyte; each microspore may develop into a male gametophyte.

3 The female and male

Gametophyte Gametophyte

Megaspore

Microspore

Meiosis

DIPLOID (2n) SPOROPHYTE GENERATION Microsporocyte

Megasporocyte Megasporangium

Archegonium

Antheridium

HAPLOID (n) GAMETOPHYTE GENERATION

1 Special cells of the

sporophyte undergo meiosis to form megaspores and microspores.

gametophytes produce gametes (eggs and sperm cells) by mitosis.

Egg

Sperm

Fertilization 4 Two haploid gametes

fuse to form a zygote.

Zygote

Microsporangium Embryo Sporophyte

5 The zygote develops

by mitosis into the sporophyte.

FIGURE 27-19 The basic life cycle of heterosporous plants Two types of spores, microspores and megaspores, are produced during the life cycle of heterosporous plants.

respective spore walls, using stored food provided by the sporophyte. As a result, and unlike the gametophytes of other seedless vascular plants, the male and female gametophytes are not truly free-living. Fertilization is followed by the development of a new sporophyte. Heterospory evolved several times during the history of land plants. It was a significant development in plant evolution because it was the forerunner of the evolution of seeds. Heterospory characterizes the two most successful groups of plants existing today, the gymnosperms and the flowering plants, both of which produce seeds.

Seedless vascular plants are used for experimental studies Botanists use many seedless vascular plants as experimental models to study certain aspects of plant biology, such as physiology, growth, and development. Ferns and other seedless vascular plants are useful in studying how apical meristems give rise to plant tissues. An apical meristem is the area at the tip (apex) of a root or shoot where growth—cell division, elongation, and differentiation—occurs. Ferns and other seedless vascular plants have a single large apical cell located at the center tip of the apical

meristem. This apical cell is the source, by mitosis, of all the cells that eventually make up the root or shoot. The apical cell divides in an orderly fashion, and the smaller daughter cells produced by the apical cell, in turn, divide and give rise to different parts of the root or shoot. It is possible to trace mature cells in the root or shoot back to their origin from the single apical cell. Ferns are interesting research plants for studies in genetics because they are polyploids and have multiple sets of chromosomes. (Many ferns have hundreds of chromosomes.) However, gene expression in ferns is exactly what one would expect of a diploid plant. Apparently, genes in the extra sets of chromosomes are gradually silenced and therefore not expressed.

Seedless vascular plants arose more than 420 mya Currently, the oldest known megafossils of early vascular plants are from Silurian (420 mya) deposits in Europe. (Plant megafossils are fossilized roots, stems, leaves, and reproductive structures.) Megafossils of several kinds of small, seedless vascular plants also occur in Silurian deposits in Bolivia, Australia, and northwestern China. Microscopic spores of early vascular plants appear in the

Sporangium

Upright stem

Rhizome

Rhizoids

FIGURE 27-20 Reconstruction of Rhynia gwynne-vaughanii

FIGURE 27-21 Reconstruction of Aglaophyton major

This leafless plant, one of Earth’s earliest vascular plants, is now extinct. It grew about 18 cm (7 in) tall. (Redrawn from Edwards, D., “Evidence for the Sporophytic Status of the Lower Devonian Plant Rhynia gwynne-vaughanii,” Review of Palaeobotany and Palynology, Vol. 29, 1980.)

Recent evidence indicates that this plant, although superficially similar to other early vascular plants, lacked conducting tissues that are characteristic of vascular plants. For this reason, it has been reclassified into a new genus and is no longer considered a rhyniophyte. (Redrawn from Mauseth, J. D., Botany: An Introduction to Plant Biology, 2nd ed., Saunders College Publishing, Philadelphia, 1995.)

fossil record earlier than megafossils, suggesting that even older megafossils of simple vascular plants may be discovered. Botanists assign the oldest vascular plants to phylum Rhyniophyta, which, according to the fossil record, arose some 420 mya and became extinct about 380 mya. The rhyniophytes are so named because many fossils of these extinct plants were found in fossil beds near Rhynie, Scotland. Rhynia gwynne-vaughanii is an example of an early vascular plant that superficially resembled whisk ferns in that it consisted of leafless upright stems that branched dichotomously from an underground rhizome (FIG. 27-20). Rhynia lacked roots, although it had absorptive rhizoids. Sporangia formed at the ends of short branches. The internal structure of its rhizome contained a central core of xylem cells for conducting water and minerals. For many years, botanists considered Rhynia major, a plant that grew about 50 cm (20 in) tall and probably lived in marshes, a classic example of a rhyniophyte. Fossils indicate that this plant had rhizoids, dichotomously branching rhizomes, and upright stems that terminated in sporangia. However, recent microscopic studies of fossil rhizomes indicate that the central core of tissue lacked the xylem cells characteristic of vascular plants. For that reason, R. major was reclassified into a new genus, Aglaophyton, and is no longer considered a rhyniophyte (FIG. 27-21). Science is an ongoing enterprise, and over time, existing knowledge is re-evaluated in

light of newly discovered evidence. Aglaophyton major is an excellent example of the self-correcting nature of science, which is in a perpetually dynamic state and changes in response to newly available techniques and data. The oldest known megafossils of fernlike trees, discovered in New York, are dated at 380 million years old. These trees, which were about 8 m (26 feet) tall, had vascular tissues but no leaves and a minimal root system. They grew vertically and reproduced by forming spores. The trunks were discovered in the late 1800s (and named Eospermatopteris), but it wasn’t until 2007 that a fossil of an entire tree was pieced together. The treetops had previously been found separate from the trunks and named Wattieza.

Review ■ ■ ■

■ ■

What adaptations do ferns have that both algae and bryophytes lack? How does one distinguish between megaphylls and microphylls? Which of the following are parts of the sporophyte generation in ferns: frond, sperm cells, egg cell, roots, sorus, sporangium, spores, prothallus, rhizome, antheridium, archegonium, and zygote? Why are whisk ferns and horsetails now classified as ferns? How does heterospory modify the plant life cycle?

■ ■

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

27.1 (page 562) 1 Discuss some environmental challenges of living on land and describe how several plant adaptations meet these challenges. The colonization of land by plants required the evolution of many anatomical, physiological, and reproductive adaptations. Plants have a waxy cuticle to protect against water loss and stomata for gas exchange needed for photosynthesis. ■ Plant life cycles have an alternation of generations in which they spend part of their life cycle in a haploid gametophyte generation and part in a diploid sporophyte generation. The gametophyte plant produces gametes by mitosis. During fertilization these gametes fuse to form a zygote, the first stage of the sporophyte generation. The zygote develops into a multicellular embryo that the gametophyte protects and nourishes. The mature sporophyte plant develops from the embryo and produces sporogenous cells (spore mother cells). These undergo meiosis to form spores, the first stage in the gametophyte generation. ■ Most plants have multicellular gametangia with a protective jacket of sterile cells surrounding the gametes. Antheridia are gametangia that produce sperm cells, and archegonia are gametangia that produce eggs. ■ Ferns and other vascular plants have xylem to conduct water and dissolved minerals and phloem to conduct dissolved sugar. 2 Name the green algal group from which plants are hypothesized to have descended and describe supporting evidence. ■ Plants probably arose from a group of green algae called charophytes. This conclusion is based in part on molecular comparisons of DNA and RNA sequences, which show the closest match between charophytes and plants. ■

an egg cell in the archegonium. The zygote grows into an embryo that develops into a moss sporophyte, which is attached to the gametophyte. Meiosis occurs within the capsule of the sporophyte to produce spores. When a spore germinates, it grows into a protonema that forms buds that develop into gametophytes. Watch the life cycles of the mosses and liverworts by clicking on the figures in CengageNOW.

27.3 (page 571) 6 Discuss the features that distinguish seedless vascular plants from algae and bryophytes. ■ Seedless vascular plants have several adaptations that algae and bryophytes lack, including vascular tissues and a dominant sporophyte generation. As in bryophytes, reproduction in seedless vascular plants depends on water as a transport medium for motile sperm cells. 7 Name and briefly describe the two phyla of seedless vascular plants. ■ Sporophytes of club mosses (phylum Lycopodiophyta) consist of roots, rhizomes, erect branches, and leaves that are microphylls. Ferns (phylum Pteridophyta) are the largest and most diverse ■ group of seedless vascular plants. The fern sporophyte consists of a rhizome that bears fronds and true roots. Phylum Pteridophyta also includes whisk ferns and horsetails. Sporophytes of whisk ferns have dichotomously branching rhizomes and erect stems; they lack true roots and leaves. Horsetail sporophytes have roots, rhizomes, aerial stems that are hollow and jointed, and leaves that are reduced megaphylls.

Explore plant evolution by clicking on the figure in CengageNOW.

27.2 (page 565) 3 Summarize the features that distinguish bryophytes from other plants. Unlike other land plants, bryophytes are nonvascular and lack xylem and phloem. Bryophytes are the only plants with a dominant gametophyte generation. Their sporophytes remain permanently attached and nutritionally dependent on the gametophytes. 4 Name and briefly describe the three phyla of bryophytes. ■ Mosses (phylum Bryophyta) have gametophytes that are green plants that grow from a filamentous protonema. ■ Many liverworts (phylum Hepatophyta) have gametophytes that are flattened, lobelike thalli; others are leafy. Hornworts (phylum Anthocerophyta) have thalloid gametophytes. ■ ■

8 Describe the life cycle of ferns and compare their sporophyte and gametophyte generations. ■ Fern sporophytes have roots, rhizomes, and leaves that are megaphylls. Their leaves, or fronds, bear sporangia in clusters called sori. Meiosis in sporangia produces haploid spores. The fern gametophyte, called a prothallus, develops from a haploid spore and bears both archegonia and antheridia. Watch the life cycle of the ferns by clicking on the figure in CengageNOW.

9 Compare the generalized life cycles of homosporous and hetero-

5 Describe the life cycle of mosses and compare their gametophyte and sporophyte generations. ■ The green moss gametophyte bears archegonia and/or antheridia at the top of the plant. During fertilization, a sperm cell fuses with

sporous plants. ■ Homospory, the production of one kind of spore, is characteristic of bryophytes, most club mosses, and most ferns, including whisk ferns and horsetails. In homospory, spores give rise to gametophyte plants that produce both egg cells and sperm cells. ■ Heterospory, the production of two kinds of spores (microspores and megaspores), occurs in certain club mosses, certain ferns, and all seed plants. Microspores give rise to male gametophytes that produce sperm cells. Megaspores give rise to female gametophytes that produce eggs. The evolution of heterospory was an essential step in the evolution of seeds.

T E S T YO U R U N D E R S TA N D I N G 1. Plants probably descended from a group of green algae called (a) rhyniophytes (b) Calamites (c) epiphytes (d) charophytes (e) club mosses 2. Which of the following is not a characteristic of plants? (a) cuticle (b) unicellular gametangia (c) stomata (d) multicellular embryo (e) alternation of generations 3. In plant life cycles (a) the first products of meiosis are gametes (b) spores are part of the diploid sporophyte generation (c) the embryo gives rise to a zygote (d) the first stage in the diploid sporophyte generation is the zygote (e) the first stage in the haploid gametophyte generation is the prothallus 4. The bryophytes (a) include mosses, liverworts, and hornworts (b) include whisk ferns, horsetails, and club mosses (c) are small plants that lack a vascular system (d) a and c (e) b and c 5. The waxy layer that covers aerial parts of plants is the (a) cuticle (b) archegonium (c) protonema (d) stoma (e) thallus 6. A strengthening compound found in cell walls of vascular plants is (a) xanthophyll (b) lignin (c) cutin (d) cellulose (e) carotenoid 7. Stomata (a) help prevent desiccation of plant tissues (b) transport water and minerals through plant tissues (c) allow gas ex-

change for photosynthesis (d) strengthen cell walls (e) produce male gametes 8. The green, gametangia-bearing moss plant (a) is the haploid gametophyte generation (b) is the diploid sporophyte generation (c) is called a protonema (d) contains cells with single large chloroplasts (e) b and c 9. _____ is a leaf that arose from a branch system. (a) An antheridium (b) A microphyll (c) A megaphyll (d) A sorus (e) A microspore 10. These plants have vascularized stems but lack true roots and leaves. (a) mosses (b) club mosses (c) horsetails (d) whisk ferns (e) hornworts 11. These plants have hollow, jointed stems that are impregnated with silica. (a) mosses (b) club mosses (c) horsetails (d) whisk ferns (e) hornworts 12. Which of the following statements about ferns is not true? (a) ferns have motile sperm cells that swim through water to the egg-containing archegonium (b) ferns are vascular plants (c) ferns are the most economically important group of bryophytes (d) the fern sporophyte consists of a rhizome, roots, and fronds (e) the diversity of ferns is greatest in the tropics

CRITICAL THINKING 1. EVOLUTION LINK. How may the following trends in plant evolution be adaptive to living on land? a. dependence on water for fertilization ¡ no need for water as a transport medium b. homospory ¡ heterospory 2. ANALYZING DATA. According to the cladogram in Figure 27-5, which plants evolved first: nonvascular bryophytes, seedless vascular plants, or seed plants? 3. EVOLUTION LINK. Where would you position the rhyniophytes on Figure 27-13? Would the line for rhyniophytes extend to the tips of the rest of the cladogram? Why or why not? 4. Label the following figures. Refer to Figs. 27-7 and 27-16 to check your answers.

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Seed Plants

© Peggy Greb/U.S. Department of Agriculture/Photo Researchers, Inc.

28

KEY CONCEPTS

Hairy vetch seed pods. The seeds are developing within each pod. Hairy vetch (Vicia villosa) is native to Europe and Asia but has become naturalized in the United States.

C

hapter 27 focused on plants that reproduce by means of spores, haploid cells that disperse and germinate to produce gametophytes.

28.1 Seed plants, which produce young sporophytes enclosed within seeds, include gymnosperms and angiosperms.

Although gymnosperms and angiosperms also produce spores, their pri-

28.2 Gymnosperms produce exposed ovules that, follow-

of an embryonic sporophyte, nutritive tissue, and a protective coat. Seeds

ing fertilization, develop into seeds that are usually borne in cones on the sporophytes. Conifers are the most diverse and numerous of the four living gymnosperm phyla.

mary means of reproduction and dispersal is by seeds, which represent an important adaptation for life on land (see photograph). Each seed consists develop from the fertilized egg cell, the female gametophyte, and its associated tissues. The two groups of seed plants, gymnosperms and angio-

28.3 Angiosperms produce ovules enclosed within car-

sperms (flowering plants), exhibit the greatest evolutionary complexity of

pels; following fertilization, seeds develop from the ovules, and the ovaries of carpels become fruits. Angiosperms dominate the land and exhibit great diversity in both vegetative and reproductive structures.

land plants and are the dominant plants in most terrestrial environments.

28.4 Gymnosperms evolved from ancestral seedless vascular plants; angiosperms evolved from ancestral gymnosperms, possibly an ancient conifer.

Seeds are reproductively superior to spores for several reasons. First, a seed is further along in its development before it is released to survive on its own: a seed contains a multicellular young plant with embryonic root, stem, and one or more leaves already formed, whereas a spore is a single cell. Second, a seed contains an abundant food supply. After germination, food stored in the seed nourishes the plant embryo until it becomes self-sufficient. Because a spore is a single cell, few food reserves exist for the plant that develops from a spore. Third, a seed is protected by a multicellular seed coat that is very thick and hard in some plants, as, for example, in lima beans. Like spores, seeds live for extended periods

at reduced rates of metabolism and germinate when conditions become favorable.

In this chapter we present the diversity in seed plants—both gymnosperms and angiosperms—followed by an examination of

Seeds and seed plants are intimately connected with the

what we know about how seed plants evolved.

development of human civilization. From prehistoric times, early humans collected and used seeds for food. Seeds often contain a concentrated source of proteins, oils, carbohydrates, and vitamins, which are nourishing for humans as well as for germinating plants. Seeds are easy to store (if kept dry), so humans can collect them

28.1 AN INTRODUCTION TO SEED PLANTS

during times of plenty to save for times of need. Few other foods

■ ■ LEARNING OBJECTIVE

are stored as conveniently or for as long. Although flowering plants

1

produce most seeds that humans consume, the seeds of certain gymnosperms—the piñon pine, for example—are edible. Embryonic leaves

Seed coat— parental sporophyte tissue

Embryo— daughter sporophyte (diploid)

Embryonic root

Haploid gametophyte (food supply)

(a) Gymnosperm seed. Longitudinal section through a pine seed.

Compare the features of gymnosperms and angiosperms.

Like the bryophytes and seedless vascular plants introduced in Chapter 27, seed plants have life cycles with an alternation of generations; they spend part of their lives in the multicellular diploid sporophyte stage and part in the multicellular haploid gametophyte stage. The sporophyte generation is the dominant stage in seed plants, and the gametophyte generation is significantly reduced in size and entirely dependent on the sporophyte generation. Unlike the bryophytes and ferns, seed plants do not have freeliving gametophytes. Instead, the female gametophyte is attached to and nutritionally dependent on the sporophyte generation. In Chapter 27, you learned that some seedless vascular plants are heterosporous. (Figure 27-19 shows a generalized life cycle for heterosporous plants.) However, all seed plants are heterosporous and produce two types of spores: microspores and megaspores. In fact, heterospory is a requirement of seed production. Seed plants produce ovules, each of which is a megasporangium surrounded by integuments, layers of sporophyte tissue that enclose the megasporangium. After fertilization takes place, the ovule develops into a seed, and the integuments develop into the seed coat (FIG. 28-1a). Botanists divide seed plants into two groups based on whether or not an ovary wall surrounds their ovules (an ovary is a structure

Gene Cox / Photo Researchers, Inc.

Ovule (future seed) surrounded by integument (future seed coat)

(b) Gymnosperm cone. Longitudinal section through a female pine cone, showing the ovules (which develop into seeds) borne on scales. Note the absence of an ovary wall.

FIGURE 28-1 Seeds

Fruit (ovary wall) Seed

(c) Angiosperm fruit. Longitudinal section through an avocado fruit, showing the seed surrounded by ovary tissue of the maternal sporophyte.

C Squared Studios /Photodisc /Getty Images

Scale

that contains one or more ovules). TABLE 28-1 A Comparison of Gymnosperms and Angiosperms The two groups of seed plants are the Characteristic Gymnosperms Angiosperms gymnosperms and the angiosperms (TABLE 28-1). The word gymnosperm is derived from the Greek for “naked seed.” Gymnosperms produce seeds that are totally exposed or borne on the scales of cones (FIG. 28-1b). In other words, an ovary wall does not surround the ovules of gymnosperms. Pine, spruce, fir, hemlock, and ginkgo are examples of gymnosperms. Growth habit Woody trees and shrubs Woody or herbaceous The term angiosperm is derived Conducting cells in xylem Tracheids Vessel elements and tracheids from the Greek expression that means Reproductive structures Cones (usually) Flowers “seed enclosed in a vessel or case.” AnPollen grain transfer Wind (usually) Animals or wind giosperms are flowering plants that Fertilization Egg and sperm ¡ zygote Double fertilization: egg and produce their seeds within a fruit (a sperm ¡ zygote; two polar nuclei and sperm ¡ endosperm mature ovary) (FIG. 28-1c). Thus, the Seeds Exposed or borne on scales Enclosed within fruit derived ovules of angiosperms are protected. of cones from ovary Flowering plants, which are extremely Nutritive material Female gametophyte Endosperm diverse, include corn, oaks, water lilNumber of species About 840 More than 300,000 ies, cacti, apples, grasses, palms, and Geographic distribution Worldwide Worldwide buttercups. Both gymnosperms and flowering plants have vascular tissues: xylem, phyta (the ginkgoes) and Cycadophyta (the cycads), are evolutionfor conducting water and dissolved minerals (inorganic nutrients), ary remnants of groups that were more significant in the past. The and phloem, for conducting dissolved sugar. fourth phylum, Gnetophyta (gnetophytes), is a collection of some unusual plants that share certain traits not found in other gymnoReview sperms; until recently, the gnetophytes were thought to be more ■ What is an ovule? closely related to flowering plants than were other gymnosperm ■ How do gymnosperm and angiosperm seeds differ? clades. Current evidence, however, suggests that gnetophytes are probably most closely related to conifers (see Fig. 28-2).

28.2 GYMNOSPERMS ■ ■ LEARNING OBJECTIVES 2 3 4

Trace the steps in the life cycle of a pine and compare its sporophyte and gametophyte generations. Summarize the features that distinguish gymnosperms from bryophytes and ferns. Name and briefly describe the four phyla of gymnosperms.

The gymnosperms include some of the most interesting plants. For example, a giant sequoia (Sequoiadendron giganteum) known as the General Sherman Tree, in Sequoia National Park in California, is one of the world’s most massive organisms. It is 82 m (267 ft) tall and has a girth of 23.7 m (77 ft) measured 1.5 m (5 ft) above ground level. Another gymnosperm, a coast redwood (Sequoia sempervirens) nicknamed “Hyperion” after a giant in Greek mythology, is possibly the world’s tallest tree, measuring 117 m (379 ft) in 2007. Botanists using tree-ring analysis determined that one of Earth’s oldest living organisms, a bristlecone pine (Pinus longaeva) in the White Mountains of California, is about 5,000 years old. Gymnosperms are usually classified into four phyla, which branch from a single evolutionary line (FIG. 28-2). Numbering 630 species, the largest phylum of gymnosperms is Coniferophyta, commonly called conifers. Two phyla of gymnosperms, Ginkgo-

Conifers are woody plants that produce seeds in cones The conifers (phylum Coniferophyta), which include pines, spruces, hemlocks, and firs, comprise the most familiar group of gymnosperms (FIG. 28-3a). These 630 species of woody trees or shrubs produce annual additions of secondary tissues (wood and bark); there are no herbaceous (nonwoody) conifers. The wood (secondary xylem) consists of tracheids, which are long, tapering cells with pits through which water and dissolved minerals move from one cell to another. Many conifers produce resin, a viscous, clear or translucent substance consisting of several organic compounds that may protect the plant from attack by fungi or insects. The resin collects in resin ducts, tubelike cavities that extend throughout the roots, stems, and leaves. Cells lining the resin ducts produce and secrete resin. Conifers generally have leaves called needles that are long, narrow, tough, and leathery (FIG. 28-3b). Most pines bear clusters of two to five needles, depending on the species. In a few conifers, such as American arborvitae, the leaves are scalelike and cover the stem (FIG. 28-3c). Most conifers are evergreen and bear their leaves throughout the year. Only a few, such as the dawn redwood, larch, and bald cypress, are deciduous and shed their needles at the end of each growing season.

FIGURE 28-2 Gymnosperm and angiosperm evolution

Most conifers are monoecious: they have separate male and female reproductive parts in different locations on the same plant. These reproductive parts are generally borne in strobili (commonly called cones), hence the name conifer, which means “cone-bearing.” Conifers occupy extensive areas, ranging from the Arctic to the tropics, and are the dominant vegetation in the forested regions of Alaska, Canada, northern Europe, and Siberia. In addition, they are important in the Southern Hemisphere, particularly in wet, mountainous areas of temperate and tropical regions in South America, Australia, New Zealand, and Malaysia. Southwestern China, with more than 60 species of conifers, has the greatest regional diversity of conifer species in the world. California, New Caledonia (an island east of Australia), southeastern China, and Japan also have considerable diversity of conifer species. Ecologically, conifers contribute food and shelter to animals and other organisms, and their roots hold the soil in place and help prevent soil erosion. Humans use conifers for their wood (for building materials as well as paper products), medicinal value (such as the anticancer drug Taxol from the Pacific yew), turpentine, and resins. Because of their attractive appearance, conifers such as firs, spruces, pines, and cedars are grown for landscape design and decorative holiday trees and wreaths.

This cladogram shows a current hypothesis of phylogenetic relationships among living seed plants, based on structural evidence, molecular comparisons, and fossils. Relationships among extant gymnosperm clades and angiosperms remain controversial. The arrangement of the phyla shown here may change as future analyses help clarify relationships.

Pines represent a typical conifer life cycle The genus Pinus, by far the largest genus in the conifers, consists of about 100 species. A pine tree is a mature

KEY POINT

Seed plants include four gymnosperm phyla and one phylum of flowering plants (angiosperms). Seed plants

Angiosperms

Gnetophytes

Conifers

Ginkgoes

Cycads

Outgroup: Ferns

Gymnosperms

Evolution of flowers

Evolution of seeds

Michael P. Gadomski /Photo Researchers, Inc.

Angiosperms

Gnetophytes

Conifers

Ginkgoes

Cycads

Outgroup: Ferns

Gymnosperms

(b) In white pine (Pinus strobus), leaves are long, slender needles that occur in clusters of five.

(a) White fir (Abies concolor). Photographed in Milford, Pennsylvania, at the historic home of Gifford Pinchot, the first Chief Forester of the U.S. Forest Service (under T. R. Roosevelt).

FIGURE 28-3 Conifers

(c) In American arborvitae (Thuja occidentalis), leaves are small and scalelike.

sporophyte (FIGURE 28-4). Pine is heterosporous and therefore produces microspores and megaspores in separate cones. Male cones, usually 1 cm or less in length, are smaller than female cones and are generally produced on the lower branches each spring (FIG. 28-5). The more familiar, woody female cones, which are on the tree year-round, are usually found on the upper branches of the tree and bear seeds after reproduction. Female cones vary considerably in size. The sugar pine (P. lambertiana) that grows in California produces the world’s longest female cones, which reach lengths of 60 cm (2 ft). Each male cone, also called a pollen cone, consists of sporophylls, leaflike scales that bear sporangia on the underside. At the base of each sporophyll are two microsporangia, which contain numerous microsporocytes, also called microspore mother cells. In

KEY POINT

Figure 28-4, each microsporocyte undergoes meiosis to form four haploid microspores. Microspores then develop into extremely reduced male gametophytes. Each immature male gametophyte, also called a pollen grain, consists of four cells, two of which—a generative cell and a tube cell—are involved in reproduction. The other two cells soon degenerate. Two large air sacs on each pollen grain provide buoyancy for wind dissemination. Male cones shed pollen grains in great numbers, and wind currents carry some to the immature female cones. Many botanists think that the female cones (also called seed cones) are modified branch systems. Each cone scale bears two ovules, or megasporangia, on its upper surface. Within each megasporangium, meiosis of a megasporocyte, or megaspore mother cell, produces four haploid megaspores. One of these di-

Gymnosperms, like all seed plants, are heterosporous.

Each scale bears two microsporangia

Microsporangium Microspores, each of which develops into a pollen grain

2

Microsporocytes within male cones undergo meiosis to form microspores, each of which develops into a pollen grain.

4

Scale from a male cone

Male cone

Pollen grains are transferred to the female cone by wind.

Scale from a female cone

3

Female gametophyte

Each scale bears two ovules (megasporangia)

Megasporocytes within female cones undergo meiosis to form megaspores.

Megasporangium Megaspore Growing pollen tube

Ovule

HAPLOID (n) GAMETOPHYTE GENERATION Meiosis Immature female cone

Fertilization DIPLOID (2n) SPOROPHYTE GENERATION

5

Zygote Second sperm nucleus Papery wings

Pollen tube

Sperm nucleus united with egg nucleus

Seed coat

Male cones (pollen cones)

6

Embryo

1

The pine tree is a mature sporophyte.

Newly germinated seedling

Female gametophyte (nutritive tissue)

Pollen tube transports sperm cells, one of which fertilizes the egg.

5

Two seeds on the upper surface of the scale

FIGURE 28-4 Animated Life cycle of pine One major advantage of gymnosperms over the seedless vascular plants is the production of wind-borne pollen grains.

Embryo in seed develops from zygote.

Mature female cone (seed cone)

Walt Anderson / Visuals Unlimited

Manfred Kage / Peter Arnold, Inc.

Pollen grains (immature male gametophytes)

not produce antheridia. The gametophyte generation in pines, as in all seed plants, depends totally on the sporophyte generation for nourishment. A major adaptation in the pine life cycle is elimination of the need for external water as a sperm transport medium. Instead, air currents carry pine pollen grains to female cones, and nonflagellate sperm cells move through a pollen tube to the egg. Pine and other conifers are plants whose reproduction is totally adapted for life on land.

50 μm

Cycads have seed cones and compound leaves Cycads (phylum Cycadophyta) were very important during the Triassic period, which began about 251 million years ago (mya) and is sometimes referred to as the “Age of Cycads.” Most species are now exFIGURE 28-5 Male and female cones in lodgepole pine (Pinus contorta) tinct, and the few surviving cycads, about Mature woody female cones (top) have opened to shed their seeds. Clusters of male cones (bottom) pro140 species, are tropical and subtropiduce copious amounts of pollen grains in the spring. (Inset) Each pollen grain develops from a microspore. cal plants with stout, trunklike stems and compound leaves that resemble those of palms or tree ferns (FIG. 28-6). Many cycads are endangered, privides mitotically, developing into the female gametophyte, which marily because they are popular as ornamentals and are gathered produces an egg within each of several archegonia. The other three from the wild and sold to collectors. In the United States, cycads megaspores are nonfunctional and soon degenerate. can be found in Florida, Texas, and California. When the ovule is ready to receive pollen, it produces a sticky Cycad reproduction is similar to that in pines except that cydroplet at the opening where the pollen grains land. Pollination, cads are dioecious and therefore have seed cones on female plants the transfer of pollen to the female cones, occurs in the spring for and pollen cones on male plants. Their seed structure is most like a week or 10 days, after which the pollen cones wither and drop off that of the earliest seeds found in the fossil record. Cycads have the tree. One of the many pollen grains that adhere to the sticky also retained motile sperm cells, each of which has many hairlike female cone grows a pollen tube, an outgrowth that digests its way flagella. Motile sperm cells are a vestige retained from the ancesthrough the megasporangium to the egg within the archegonium. tors of cycads, in which sperm cells swam from antheridia to archeThe germinated pollen grain with its pollen tube is the mature male gonia. In cycads, specialized insects (almost exclusively beetles) gametophyte. carry pollen grains to the female plants and their cones. Following Ultimately, two nonmotile (nonflagellate) sperm cells form transfer to the female cone, the pollen grain germinates and grows within the germinated pollen grain. When it reaches the female a pollen tube. The sperm cells are released into this tube and swim gametophyte, the pollen tube discharges the two sperm cells near to the egg. the egg. One of these sperm cells fuses with the egg, in the process of fertilization, to form a zygote, or fertilized egg, which subsequently grows into a young pine embryo in the seed. The other Ginkgo biloba is the only sperm cell degenerates. living species in its phylum The developing embryo consists of an embryonic root and Ginkgo (phylum Ginkgophyta) is represented by a single extant an embryonic shoot with several cotyledons (embryonic leaves). (living) species, the maidenhair tree, Ginkgo biloba (FIG. 28-7). Haploid female gametophyte tissue surrounds the embryo and beThis native of eastern China grew in the wild in only two locations, comes the nutritive tissue in the mature pine seed. A tough, protecalthough people had cultivated it for its edible seeds in China and tive seed coat derived from the integuments encloses the embryo Japan for centuries. Ginkgo is the oldest genus (and species) of exand nutritive tissue. The seed coat forms a thin, papery wing at tant trees. Fossil ginkgoes 200 million years old have been discovone end that enables dispersal by air currents. Some seeds remain ered that are strikingly similar to the modern ginkgo. within the female cones for several years before being shed. People often plant ginkgo in North America and Europe today, In the pine life cycle, the sporophyte generation is dominant, particularly in parks and along city streets, because it is hardy and and the gametophyte generation is restricted in size to microscopic somewhat resistant to air pollution. Its leaves are deciduous and structures in the cones. Although the female gametophyte proturn a beautiful yellow before being shed in the fall. duces archegonia, the male gametophyte is so reduced that it does

Gnetophytes include three unusual genera

Female strobilus (seed cone)

(a) A female Florida arrowroot (Zamia

(b) This female cycad (Encephalartos transvenosus) in South Africa has a trunk that reaches a height of about 9 m (30 ft) and resembles a palm. Note its immense seed cones that grow up to 0.8 m (30 in) in length.

Marion Lobstein

Joseph Malcolm Smith/Photo Researchers, Inc.

integrifolia) produces seed cones. This plant The gnetophytes (nee¿ toe phites) is the only cycad native to the United States. (phylum Gnetophyta) consist of Like Zamia, most cycads are short plants, about 70 species in three diverse less than 2 m (6.5 ft) tall. and obscure genera (Gnetum, Ephedra, and Welwitschia). Gnetophytes FIGURE 28-6 Cycads share certain features that make them unique among the gymnosperms. For example, gnetophytes have more efficient waterconducting cells, called vessel elements, in their xylem (see Chapter 33). Flowering plants also have vessel elements in their xylem, but of the gymnosperms, only the gnetophytes do. In addition, the cone clusters that some gnetophytes produce resemble flower clusters, and certain details in their life cycles resemble those of flowering plants. Despite these similarities, most botanists now think that

Walter H. Hodge / Peter Arnold, Inc.

Like cycads, ginkgo is dioecious, with separate male and female trees. It has flagellate sperm cells—an evolutionary vestige that is not required because ginkgo produces airborne pollen grains. Ginkgo seeds are completely exposed rather than contained within cones. Male trees are usually planted because the female trees bear seeds whose fleshy outer coverings give off a foul odor that smells like rancid butter. In China and Japan, where people eat the seeds, the female trees are more common. Ginkgo has been an important medicinal plant for centuries and is still a common herbal remedy today. Extracts from the leaves may enhance neurological functioning by increasing blood flow to the brain, although several recent studies have concluded that ginkgo does not improve memory in elderly people.

(a) Close-up of a branch from a female ginkgo, showing the exposed seeds and the distinctive, fanshaped leaves.

FIGURE 28-7 Ginkgo, or maidenhair tree

(b) Ginkgo (Ginkgo biloba) tree in a formal garden in northern England.

Review

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What is the dominant generation in the pine life cycle? How does pollination occur in gymnosperms? What features distinguish gymnosperms from other plants? What are the four groups of gymnosperms?

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What features distinguish cycads from ginkgo? from gnetophytes?

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28.3 FLOWERING PLANTS ■ ■ LEARNING OBJECTIVES 5

Summarize the features that distinguish flowering plants from other plants.

6 Briefly explain the life cycle of a flowering plant and describe double 7 8

fertilization. Contrast monocots and eudicots, the two largest classes of flowering plants. Discuss the evolutionary adaptations of flowering plants.

Flowering plants, or angiosperms (phylum Anthophyta), are the most successful plants today, surpassing even the gymnosperms in importance. They have adapted to almost every habitat and, with at least 300,000 species, are Earth’s dominant plants. Flowering

David Cavagnaro

(b) A male joint fir (Ephedra) has pollen cones clustered at the nodes. In the 19th century, European pioneers used species native to the American Southwest to make a beverage, Mormon tea.

Robert and Linda Mitchell

Dr. John D. Cunningham/ Visuals Unlimited, Inc.

gnetophytes represent evolutionary diversity in gymnosperms and are not in a direct line to flowering plants (see Fig. 28-2). The genus Gnetum contains tropical vines, shrubs, and trees with broad leaves (FIG. 28-8a). Species in the genus Ephedra include many shrubs and vines that grow in deserts and other dry temperate and tropical regions. Some Ephedra species resemble horsetails in that they have jointed green stems with tiny leaves (FIG. 28-8b). Commonly called joint fir, Ephedra has been used medicinally for centuries. An Asiatic Ephedra is the source of ephedrine, which stimulates the heart and raises blood pressure. At one time, ephedrine was commonly sold over (a) The leaves of Gnetum gnemon the counter in weight-control medicaresemble those of flowering plants. Note tions and herbal energy boosters; several the exposed seeds. The species is native deaths were reported from chronic use or to southern Asia and the Maldives. overdose of products containing ephedrine, so its use has been restricted. The third gnetophyte genus, Welwitschia, contains a single species found in deserts of southwestern Africa (FIG. 28-8c). Most of Welwitschia’s body—a long taproot—grows underground. Its short, wide stem forms a shallow disc, up to 0.9 m (3 ft) in diameter, from which two ribbonlike leaves extend. These two leaves continue to grow from the stem throughout the plant’s life, but their ends are usually broken and torn by the wind, giving the appearance of numerous leaves. Each leaf grows to about 2 m (6.5 ft) in length. When Welwitschia reproduces, cones form around the edge of its disclike stem.

(c) Welwitschia mirabilis is native to deserts in southwestern Africa. It survives on moisture-laden fogs that drift inland from the ocean. Photographed in the Namib Desert, Namibia.

FIGURE 28-8 Gnetophytes

plants come in a wide variety of sizes and forms, from herbaceous violets to massive eucalyptus trees. Some flowering plants—tulips and roses, for example—have large, conspicuous flowers; others, such as grasses and oaks, produce small, inconspicuous flowers. Flowering plants are vascular plants that reproduce sexually by forming flowers and, following a unique double fertilization process, seeds within fruits. The fruit protects the developing seeds and often aids in their dispersal (see Chapter 37). Flowering plants have efficient water-conducting cells called vessel elements in their xylem and efficient sugar-conducting cells called sieve tube elements in their phloem (see Chapter 33). Flowering plants are extremely important to humans because our survival as a species literally depends on them. All our major

food crops are flowering plants; examples include rice, wheat, corn, potatoes, tomatoes, beans, apples, and citrus fruits. Woody flowering plants such as oak, cherry, and walnut provide valuable lumber. Flowering plants give us fibers, such as cotton and linen, and medicines, such as digitalis and codeine. Products as diverse as rubber, tobacco, coffee, chocolate, wine, and aromatic oils for perfumes come from flowering plants. Economic botany is the subdiscipline of botany that deals with plants of economic importance.

TABLE 28-2

Distinguishing Features of Monocots and Eudicots

Feature

Monocots

Eudicots

Flower parts Pollen grains Leaf venation Vascular bundles in stem cross section Roots Seeds Secondary growth (wood and bark)

Usually in threes One furrow or pore Usually parallel Usually scattered or more complex arrangement Fibrous root system Embryo with one cotyledon Absent

Usually in fours or fives Three furrows or pores Usually netted Arranged in a circle (ring) Taproot system Embryo with two cotyledons Often present

Monocots and eudicots are the two largest classes of flowering plants carpels (these flower parts are discussed shortly). Monocot seeds have a single cotyledon, or embryonic seed leaf; endosperm, a nutritive tissue, is usually present in the mature seed. Eudicots include oaks, roses, mustards, cacti, blueberries, and sunflowers. Eudicots are either herbaceous (such as a tomato plant) or woody (such as a hickory tree). Their leaves vary in shape but usually are broader than monocot leaves, with netted (finely branched) veins. Flower parts usually occur in fours or fives or multiples thereof (FIG. 28-9b). Two cotyledons are present in eudicot seeds, and endosperm is usually absent in the mature seed, having been absorbed by the two cotyledons during seed development.

Sexual reproduction takes place in flowers

Angiosperms

Flowers are reproductive shoots usually composed of four parts— sepals, petals, stamens, and carpels—arranged in whorls (circles)

Richard H. Gross

John Gerlach/Tom Stack & Associates

Gnetophytes

Conifers

Ginkgoes

Cycads

Outgroup: Ferns

Phylum Anthophyta is divided into several classes with only a few members each and two very large classes: the monocots (class Monocotyledones) and the eudicots (class Eudicotyledones). The smaller classes will be discussed later in the chapter in the context of their evolutionary significance. For now, we restrict our discussion of flowering plants to the monocots and eudicots, which collectively represent about 97% of all flowering plant species. Eudicots are more diverse and include many more species (at least 200,000) than the monocots (at least 90,000). TABLE 28-2 provides a comparison of some of the general features of the two classes. Monocots include palms, grasses, orchids, irises, onions, and lilies. Monocots are mostly herbaceous plants with long, narrow leaves that have parallel veins (the main leaf veins run parallel to one another). The parts of monocot flowers usually occur in threes (FIG. 28-9a). For example, a flower may have three sepals, three petals, six stamens, and a compound pistil consisting of three fused

(a) Monocot. Trillium erectum, like most monocots, has floral parts in threes. Note the three green sepals, three red petals, six stamens, and three stigmas (the compound pistil consists of three fused carpels).

FIGURE 28-9 Flowering plants

(b) Eudicot. Most eudicots, such as this Tacitus, have floral parts in fours or fives. Note the five petals, 10 stamens, and five separate pistils. Five sepals are also present but barely visible against the background.

Darwin Dale / Photo Researchers, Inc.

on the end of a flower stalk, or peduncle (FIG. 28-10). The peduncle may terminate in a single flower or a cluster of flowers known as an inflorescence. The tip of the flower stalk that bears the flower parts is known as the receptacle. All four floral parts are important in the reproductive process, but only the stamens (the “male” organs) and carpels (the

“female” organs) produce gametes. A flower that has all four parts is complete, whereas an incomplete flower lacks one or more of these four parts. A flower with both stamens and carpels is perfect, whereas an imperfect flower has stamens or carpels, but not both. Sepals, which make up the lowermost and outermost whorl on a floral shoot, are leaflike in appearance and often green (FIG. 28-11a). Sepals cover and protect the other flower parts when the flower is a bud. As the flower opens, the sepals fold back to reveal the more conspicuous petals. The collective term for all the sepals of a flower is the calyx. The whorl just above the sepals consists of petals, which are broad, flat, and thin (like sepals and leaves) but vary in shape and are frequently brightly colored. Petals attract animal pollinators to the flower (see Chapter 37). Sometimes petals are fused to form a tube (as in trumpet honeysuckle flowers) or other floral shape (as in snapdragons, whose petals form two lips). The petals of a flower are referred to collectively as the corolla. Just inside the petals is a whorl of stamens (FIG. 28-11b). Each stamen is composed of a thin stalk, called a filament, and a saclike anther, where meiosis occurs to form microspores that develop into pollen grains. Each pollen grain produces two cells surrounded by a tough outer wall. One cell eventually divides to form two male gametes, or sperm cells, and the other produces a pollen tube through which the sperm cells travel to reach the ovule. In the center of most flowers are one or more closed carpels, the “female” reproductive organs. Carpels bear ovules, which, as

(a) An Arabidopsis thaliana flower. Female floral parts

Male floral parts Pollen grain (each will produce two sperm cells)

Stigma Style PISTIL (consisting of one or more carpels)

Anther STAMEN Filament

Ovary Ovules (each producing one egg cell)

Petal

Receptacle

Sepal

Peduncle

(b) This cutaway view of an Arabidopsis flower shows the details of basic floral structure. Each flower has four sepals (two are shown), four petals (two are shown), six stamens, and one long pistil. Four of the stamens are long, and two are short (two long and two short are shown). Pollen grains develop within sacs in the anthers. In Arabidopsis, the compound pistil consists of two carpels that each contain numerous ovules.

FIGURE 28-10 Animated Floral structure

Sepals

James Mauseth/University of Texas

Petals

Stamen

Pistil

(a) The leaflike sepals of a rosebud (Rosa) enclose and protect the inner flower parts.

Marion Lobstein

FIGURE 28-11 Parts of a flower

Stigma Ovules

(b) A twinleaf (Jeffersonia diphylla) flower has eight yellow stamens. Note the simple pistil with its green ovary in the center of the flower.

Style

Ovary wall

Ovary

(a) Simple pistil. This simple pistil consists of a single carpel. Ovules

One carpel

you may recall, are structures with the potential to develop into seeds. The carpels of a flower are separate or fused into a single structure. The female part of the flower is also called a pistil (see Fig. 28-11b). A pistil may consist of a single carpel (a simple pistil) or a group of fused carpels (a compound pistil) (FIG. 28-12). Each pistil generally has three sections: a stigma, on which the pollen grain lands; a style, a necklike structure through which the pollen tube grows; and an ovary, an enlarged structure that contains one or more ovules. Each young ovule contains a female gametophyte that forms one female gamete (an egg), two polar nuclei, and several other haploid cells. After fertilization, the ovule develops into a seed; and the ovary, into a fruit.

Stigma

The life cycle of flowering plants includes double fertilization Style

Ovary wall Ovary

(b) Compound pistil. This compound pistil has two united carpels. In most flowers with single pistils, the pistils are compound, consisting of two or more fused carpels.

FIGURE 28-12 Simple and compound pistils

Flowering plants undergo alternation of generations in which the sporophyte generation is larger and nutritionally independent (FIG. 28-13). The gametophyte generation in flowering plants is microscopic and nutritionally dependent on the sporophyte. Flowering plants, like gymnosperms and certain other vascular plants, are heterosporous and produce two kinds of spores: microspores and megaspores. Sexual reproduction occurs in the flower. Each young ovule within an ovary contains a megasporocyte (megaspore mother cell) that undergoes meiosis to produce four haploid megaspores. Three of these usually disintegrate, and one divides mitotically and develops into a mature female gametophyte, also called an embryo sac. Embryo sacs in the vast majority of angiosperms contain seven cells with eight haploid nuclei. Six of these cells, including the egg cell, contain a single nucleus each, and a central cell has two nuclei,

KEY POINT

A significant feature of the flowering plant life cycle is double fertilization, in which one sperm cell unites with the egg, forming a zygote; the other sperm cell unites with the two polar nuclei, forming a triploid cell that gives rise to endosperm. Developing pollen tube of mature male gametophyte

Pollination

4

Each microspore develops into a pollen grain.

Microspore Tetrad of microspores 3

6

Pollen tube grows through female tissue to deposit two sperm cells in embryo sac.

Embryo sac (mature female gametophyte)

Pollen grain (immature male gametophyte)

Pollen tube

Microsporocytes within anther sacs undergo meiosis to form haploid microspores.

Polar nuclei

One megaspore Megaspore develops into embryo sac HAPLOID (n) GAMETOPHYTE (female gametophyte). 5

GENERATION

Meiosis 2

Double fertilization

DIPLOID (2n)

Megasporocyte undergoes SPOROPHYTE meiosis to form haploid GENERATION megaspores. Ovary Megasporocyte Megasporangium (ovule)

Two sperm cells

Egg nucleus

7

Double fertilization occurs, resulting in diploid zygote and triploid endosperm.

Endosperm (3n) Zygote (2n)

Fruit

Embryo Microsporocytes within microsporangia

Seed coat

Seed

Endosperm Seedling

Anther 1

Mature sporophyte of angiosperms produces flowers for reproduction.

FIGURE 28-13 Animated Life cycle of flowering plants

called polar nuclei. The egg and the central cell with two polar nuclei are directly involved in fertilization; the other five cells in the embryo sac apparently have no direct role in the fertilization process and disintegrate. As the synergids (the two cells flanking the egg) disintegrate, however, they release chemicals that may affect the direction of pollen tube growth. Each pollen sac, or microsporangium, of the anther contains numerous microsporocytes (microspore mother cells), each of which undergoes meiosis to form four haploid microspores. Every microspore develops into an immature male gametophyte, also called a pollen grain. Pollen grains are small; each consists of two cells, the tube cell and the generative cell. The anthers split open and begin to shed pollen. A variety of agents—including wind, water, insects, and other animal pollinators—transfer pollen grains to the stigma (see Chapter 37). If compatible with the stigma, the pollen grain germinates, and a pollen tube grows down the style and into the ovary. The germi-

nated pollen grain with its pollen tube is the mature male gametophyte. Next, the generative cell divides to form two nonflagellate sperm cells. The sperm cells move down the pollen tube and are discharged into the embryo sac. Both sperm cells are involved in fertilization. When the two sperm cells enter the embryo sac, both participate in fertilization. One sperm cell fuses with the egg, forming a zygote that grows by mitosis and develops into a multicellular embryo in the seed. The second sperm cell fuses with the two haploid polar nuclei of the central cell to form a triploid (3n) cell that divides by mitosis and develops into endosperm, a nutrient tissue rich in lipids, proteins, and carbohydrates that nourishes the growing embryo. This fertilization process, which involves two separate nuclear fusions, is called double fertilization and is, with two exceptions, unique to flowering plants. (Double fertilization has been reported in the gymnosperms Ephedra nevadensis and Gnetum gnemon. This process differs from double fertilization in

Holt Studies Intl./Silvestra Silva/Photo Researchers, Inc.

FIGURE 28-14 Guava fruit and seeds Animals eat fleshy fruits such as guava. The seeds are frequently swallowed whole and pass unharmed through the animals’ digestive tracts.

flowering plants in that an additional zygote, rather than endosperm, is produced. The second zygote later disintegrates.)

Seeds and fruits develop after fertilization As a result of double fertilization and subsequent growth and development, each seed contains a young plant embryo and nutritive tissue (the endosperm), both of which are surrounded by a protective seed coat. In monocots the endosperm persists and is the main source of food in the mature seed. In most eudicots the endosperm nourishes the developing embryo, which subsequently stores food in its cotyledons. As a seed develops from an ovule following fertilization, the ovary wall surrounding it enlarges dramatically and develops into a fruit. In some instances, other tissues associated with the ovary also enlarge to form the fruit (see discussion of fruits in Chapter 37). Fruits serve two purposes: to protect the developing seeds from desiccation as they grow and mature and to aid in the dispersal of seeds. For example, dandelion fruits have feathery plumes that are lifted and carried by air currents. Animals often assist in dispersing seeds found in edible fruits (FIG. 28-14). Once a seed lands in a suitable place, it may germinate and develop into a mature sporophyte that produces flowers, and the life cycle continues as described.

Flowering plants have many adaptations that account for their success The evolutionary adaptations of flowering plants account for their success in terms of their ecological dominance and their great number of species. Seed production as the primary means of re-

production and dispersal, an adaptation shared with the gymnosperms, is clearly significant and provides a definite advantage over seedless vascular plants. Closed carpels, which give rise to fruits surrounding the seeds, and the process of double fertilization with its resulting endosperm, increase the likelihood of reproductive success. The evolution of a variety of interdependencies with many types of insects, birds, and bats, which disperse pollen from one flower to another of the same species, is another reason for angiosperm success. Pollen transfer results in cross-fertilization, which mixes the genetic material and promotes genetic variation among the offspring. Several distinctive features have contributed to the success of flowering plants in addition to their highly successful reproduction involving flowers, fruits, and seeds. Recall that most flowering plants have very efficient water-conducting vessel elements in their xylem, as well as tracheids. In contrast, the xylem of almost all seedless vascular plants and gymnosperms consists exclusively of tracheids. Most flowering plants also have efficient carbohydrateconducting sieve tube elements in their phloem. Vascular plants other than flowering plants and gnetophytes lack vessel elements and sieve tube elements. The leaves of flowering plants, with their broad, expanded blades, are very efficient at absorbing light for photosynthesis. Abscission (shedding) of these leaves during cold or dry periods reduces water loss and has enabled some flowering plants to expand into habitats that would otherwise be too harsh for survival. The stems and roots of flowering plants are often modified for food or water storage, another feature that helps flowering plants survive in severe environments. Probably most crucial to the evolutionary success of flowering plants, however, is the overall adaptability of the sporophyte generation. As a group, flowering plants readily adapt to new habitats and changing environments. This adaptability is evident in the great diversity of growth forms exhibited by the various species of flowering plants. For example, the cactus is remarkably well-adapted to desert environments. Its stem stores water; its leaves (spines) have a reduced surface area available for transpiration (loss of water vapor; see Chapter 34) and may also protect against thirsty herbivorous animals; and its thick, waxy cuticle reduces water loss. In contrast, the water lily is well-adapted for wet environments, in part because it has air channels that provide adequate oxygen to stems and roots living in oxygen-deficient water and mud.

Floral structure provides insights into the evolutionary process In evolution, new structures or organs often originate by modification of previously existing structures or organs. (See Chapter 20 discussion of preadaptations.) Much evidence supports the classical interpretation that the four organs of a flower—sepals, petals, stamens, and carpels—arose from highly modified leaves. This evidence includes comparisons of the arrangement of vascular tissues in both flowers and leafy stems and of the developmental stages of floral parts and leaves. Sepals are the most leaflike of the four floral organs, and botanists generally agree that sepals are specialized leaves. Although petals of many flowering plant species are leaflike in appearance,

botanists generally view petals as modified stamens that later became sterile and leaflike. Cultivated roses and camellias provide evidence supporting this hypothesis; in some varieties the stamens have been transformed into petals, forming showy flowers with large numbers of petals. The remarkably leaflike stamens and carpels of certain tropical trees and other species support the origin of stamens and carpels from leaves or leaflike organs. Consider, for example, the carpel of Drimys, a genus of flowering trees and shrubs native to Southeast Asia, Australia, and South America. This carpel resembles a leaf that is folded inward along the midrib, thereby enclosing the ovules, and joined along the entire length of the leaf ’s margin (FIG. 28-15). The fundamental question is whether these leaflike stamens and carpels are shared ancestral characters that were conserved (retained) during the course of evolution or are highly specialized organs (that is, shared derived characters) that do not resemble early stamens and carpels. Many botanists who have studied this question have concluded that stamens and carpels are probably derived from leaves. Not all botanists accept the origin of stamens and carpels from highly modified leaves, however. As we noted in Chapter 1, uncertainty and debate are part of the scientific process, and scientists can never claim to know a final answer. During the course of more than 130 million years of angiosperm evolution, floral structure diversified as floral organs fused or became reduced in size or number. These changes led to greater complexity in floral structure in some species and to greater simplicity in other species. Interpreting the floral structures of so many different angiosperm species is sometimes difficult, but it is important because correct interpretations are essential to devising a phylogenetic classification scheme.

Review ■

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How do nonreproductive adaptations of flowering plants differ from those of gymnosperms? How does the flowering plant life cycle differ from that of the gymnosperms? What are the two major classes of flowering plants, and how can one distinguish between them? How does fertilization differ in gymnosperms and flowering plants?

28.4 THE EVOLUTION OF SEED PLANTS ■ ■ LEARNING OBJECTIVE 9 Summarize the evolution of gymnosperms from seedless vascular plants and trace the evolution of flowering plants from gymnosperms.

One group that descended from ancestral seedless vascular plants was the progymnosperms, all of which are now extinct. Progymnosperms had two derived features: leaves with branching veins (megaphylls) and woody tissue (secondary xylem) similar to that of modern gymnosperms. Progymnosperms, however, reproduced by spores, not seeds. Archaeopteris, a progymnosperm that lived

Carpel

Fused margin

Ovules

Stigma

Ovary

Style

Stigma

(a) The carpel resembles a folded leaf in which the ovules borne on its upper surface are enclosed.

(b) A cross section of the carpel, cut along the dashed line in (a).

FIGURE 28-15 Carpel of Drimys piperita

about 370 mya, is one of the earliest known trees with “modern” woody tissue (FIG. 28-16a). Fossils of several progymnosperms with reproductive structures intermediate between those of spore plants and seed plants have been discovered. For example, the evolution of microspores into pollen grains and of megasporangia into ovules (seedproducing structures) can be traced in fossil progymnosperms. Plants producing seeds appeared during the late Devonian period, more than 359 mya. The fossil record indicates that different groups of seed plants apparently arose independently several times. As mentioned previously, fossilized remains of ginkgo are found in 200-million-year-old rocks, and other groups of gymnosperms were well established by 160 mya to 100 mya. Although the gymnosperms are an ancient group, some questions persist about the exact pathways of gymnosperm evolution. The fossil record indicates that progymnosperms probably gave rise to conifers and to another group of extinct plants called seed ferns, which were seed-bearing woody plants with fernlike leaves (FIG. 28-16b). The seed ferns, in turn, probably gave rise to cycads and ginkgo, as well as to several gymnosperm groups now extinct. The origin of gnetophytes remains unclear, although molecular data indicate that they are closely related to conifers.

Our understanding of the evolution of flowering plants has made great progress in recent years Flowering plants are the most recent clade of plants to evolve. The fossil record, although incomplete, suggests that flowering plants descended from gymnosperms. By the middle of the Jurassic period, about 180 mya, several gymnosperm lines existed with some features resembling those of flowering plants. Among other traits, these derived gymnosperms possessed leaves with broad, expanded blades and the first modified seed-bearing leaves, which nearly enclose the ovules. Beetles were evidently visiting these plants, and biologists have suggested that perhaps this relationship

was the beginning of coevolution, a mutual adaptation between plants and their animal pollinators (see Chapter 37). One important task facing paleobotanists (biologists who study fossil plants) is determining which of the ancient gymnosperms are in the direct line of evolution leading to the flowering plants. Given the structural data, most botanists hypothesize that flowering plants arose only once—that is, there is only one line of evolution from the gymnosperms to the flowering plants. Based largely on structural data, many evolutionary botanists previously thought that the gnetophytes were the gymnosperm clade most closely related to flowering plants. However, new hypotheses of angiosperm origin have been proposed in recent years with advances in molecular comparisons, genetic studies (particularly as gene function relates to reproductive development), and additional fossil discoveries. Currently, many botanists think the closest extant gymnosperms related to flowering plants are probably conifers. But this hypothesis is far from certain, and the origin and early evo(a) Progymnosperm. Archaeopteris, (b) Seed fern. Emplectopteris produced lution of flowering plants continue to chalwhich existed about 370 mya, had seeds on fernlike leaves. Seed ferns some features in common with modern existed from about 360 mya to 250 mya. lenge botanists. seed plants but did not produce seeds. The oldest definitive trace of flowering plants in the fossil record consists of ovules FIGURE 28-16 Evolution of seed plants enclosed in tiny podlike fruits interpreted (a, Redrawn from Beck, C. B., “Reconstructions of Archaeopteris and Further Consideration of Its Phyas carpels in Jurassic and Lower Cretaceous logenetic Position,” American Journal of Botany, Vol. 49, 1962. b, Redrawn from Andrews, H. N., Ancient rocks some 125 million to 145 million years Plants and the World They Lived In, Comstock, New York, 1947.)

Pistils

Scars on reproductive axis

Ovule

5 mm

(a) The oldest known fossil angiosperm. This fossil of Archaefructus shows a carpel-bearing stem. Discovered in northeastern China, it is about 125 million years old.

FIGURE 28-17 Fossil angiosperms

Courtesy of David Dilcher

David Dilcher and Ge Sun

Carpel

7 mm

(b) Fossil flower. The fossilized flower of the extinct plant Archaeanthus linnenbergeri, which lived about 100 mya. The scars on the reproductive axis (receptacle) may show where stamens, petals, and sepals were originally attached but abscised (fell off). Many spirally arranged pistils were still attached at the time this flower was fossilized.

old (FIG. 28-17a). The oldest fossilized flowers are about 118 million to 120 million years old. By 90 mya, during the Cretaceous period, flowering plants had diversified and had begun to replace gymnosperms as Earth’s dominant plants. Fossils of flowering plant leaves, stems, flowers, fruits, and seeds are numerous and diverse. They outnumber fossils of gymnosperms and ferns in Late Cretaceous deposits, indicating the rapid success of flowering plants once they appeared (FIG. 28-17b). Many angiosperm species apparently arose from changes in chromosome number. (See discussion of allopolyploidy and sympatric evolution in Chapter 20.)

Flowering plants evolved from ancient gymnosperms. Basal angiosperms are early diverging clades that still survive today.

Magnoliids

Star anise

Water lilies

Amborella

Outgroup: Gymnosperms

Eudicots

Core angiosperms

Basal angiosperms

Monocots

KEY POINT

The basal angiosperms comprise three clades Beginning in 1999, a series of cladistic analyses of structural features and molecular comparisons have helped clarify relationships among the angiosperm classes. In the process, many long-held assumptions about early angiosperm lineages have been discarded. The evidence indicates that several clades of basal (early-diverging) angiosperms evolved before the divergence of core angiosperms (FIG. 28-18a). The 170 species of basal angiosperms represent groups thought to be ancestral to all other flowering plants. The oldest surviving clade of basal angiosperms is represented by a single extant species, Amborella trichopoda (FIG. 28-18b). A

Evolution of vessel elements Evolution of flowers

(b) Amborella trichopoda, a basal angiosperm.

(c) Water lily (Nymphaea), a basal angiosperm.

FIGURE 28-18 Evolution of flowering plants

Sinclair Stammers /Photo Researchers, Inc.

Robert and Ann Simpson/ Visuals Unlimited, Inc.

Rod Planck /Photo Researchers, Inc.

Stephen McCabe, Arboretum University of California Santa Cruz

(a) One hypothesis of relationships among the flowering plants, based on fossil and molecular evidence. Amborella, water lilies, and star anise are living plants whose ancestors apparently branched off of the angiosperm family tree early. These early clades were followed by the magnoliids, the monocots, and the eudicots.

(d) Star anise (Illicium verum), a basal angiosperm.

(e) Magnolia grandiflora, a core angiosperm.

The core angiosperms comprise magnoliids, monocots, and eudicots Most angiosperm species belong to a clade of core angiosperms, which is divided into three subclades: magnoliids, monocots, and eudicots. Magnoliids include species in the magnolia, laurel, and black pepper families, as well as several related families (FIG. 28-18e). Although magnoliids were traditionally classified with the eudicots as “dicots,” molecular evidence such as DNA sequence comparisons indicates that the magnoliids are neither eudicots nor monocots. Native to tropical or warm temperate regions, magnoliids include several economically important plants, such as avocado, black pepper, nutmeg, and bay laurel.

Review What features distinguish progymnosperms from seed ferns? Describe the significant features of the oldest known fossil angiosperm. Are monocots considered basal or core angiosperms? Explain your answer.

■ ■

■

■ ■

KEY EXPERIMENT QUESTION: Hydatella sp. was incorrectly classified as a grass until recent molecular evidence indicated that it is a basal angiosperm closely related to water lilies. Does Hydatella have any reproductive features that may indicate it is one of the most ancient surviving lineages of angiosperms? EXPERIMENT: Hydatella inconspicua, a small aquatic plant, was collected at a lake in Northland, New Zealand, and its embryo sacs were carefully analyzed. © Justin Goh, School of Biological Sciences, University of Auckland, Auckland, New Zealand

shrub native to New Caledonia, an island in the South Pacific, Amborella may be the nearest living relative to the ancestor of all flowering plants. The water lilies and related families compose the second clade of basal angiosperms (FIG. 28-18c and FIG. 28-19). This clade, which contains about 70 species of aquatic or wetland herbs, may be the secondoldest surviving lineage. Star anise and relatives—the third clade of basal angiosperms—consist of about 100 species of vines, trees, and shrubs found mostly in warmer climates. Star anise is important economically because it is a source of spice used by confectioners and anise oil (FIG. 28-18d). Star anise is also used to make Tamiflu, one of the treatments for influenza.

RESULTS AND CONCLUSION: As in gymnosperms, the mother plant turns over nutrients to the seed before fertilization has occurred. This gymnosperm-like reproductive feature may represent a shared ancestral character that was retained during the transition from gymnosperms to flowering plants, thus supporting the view that Hydatella is a representative of an ancient angiosperm lineage. Alternatively, this feature may represent a shared derived character not found in earlier Hydatella ancestors. Source: Friedman, W. E. “Hydatellaceae are Water Lilies with Gymnospermous Tendencies.” Nature, Vol. 453, May 1, 2008.

FIGURE 28-19 The basal angiosperm Hydatella Hydatella inconspicua belongs to an ancient clade that consists of about 10 species native to Australia, New Zealand, and India.

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

28.1 (page 583)

■

1 Compare the features of gymnosperms and angiosperms. ■

The two groups of seed plants are the gymnosperms and the angiosperms. Gymnosperms produce seeds that are totally exposed or borne on the scales of cones; an ovary wall does not surround the ovules of gymnosperms. Angiosperms are flowering plants that produce their seeds within a fruit (a mature ovary).

28.2 (page 584) 2 Trace the steps in the life cycle of a pine and compare its sporophyte and gametophyte generations. A pine tree is a mature sporophyte; pine gametophytes are extremely small and nutritionally dependent on the sporophyte generation. Pine is heterosporous and produces microspores and megaspores in separate cones. Male cones produce microspores that develop into pollen grains ■ (immature male gametophytes) that are carried by air currents to female cones. ■

■

Female cones produce megaspores. One of each four megaspores produced by meiosis develops into a female gametophyte within an ovule (megasporangium). After pollination, the transfer of pollen to the female cones, a pollen tube grows through the megasporangium to the egg within the archegonium. After fertilization, the zygote develops into an embryo encased inside a seed adapted for wind dispersal.

3 Summarize the features that distinguish gymnosperms from bryophytes and ferns. ■ Unlike bryophytes, gymnosperms are vascular plants. Unlike bryophytes and ferns, gymnosperms produce seeds. Gymnosperms also produce wind-borne pollen grains, a feature absent in ferns and other seedless vascular plants. 4 Name and briefly describe the four phyla of gymnosperms. ■ Conifers (phylum Coniferophyta), the largest phylum of gymnosperms, are woody plants that bear needles (leaves that are usually evergreen) and produce seeds in cones. Most conifers are monoecious and have male and female reproductive parts in separate cones on the same plant. ■ Cycads (phylum Cycadophyta) are palmlike or fernlike in appearance. They are dioecious—they have male and female reproductive structures on separate plants—but reproduce with pollen and seeds in conelike structures. ■ Ginkgo biloba, the only surviving species in phylum Ginkgophyta, is a deciduous, dioecious tree. The female ginkgo produces fleshy seeds directly on branches. Gnetophytes (phylum Gnetophyta) are an obscure clade of gymno■ sperms that has a few traits associated with angiosperms.

28.3 (page 589) 5 Summarize the features that distinguish flowering plants from other plants. Flowering plants, or angiosperms (phylum Anthophyta), constitute the phylum of vascular plants that produce flowers and seeds enclosed within a fruit. They are the most diverse and most successful group of plants. ■ The flower, which may contain sepals, petals, stamens, and carpels, functions in sexual reproduction. Unlike those of gymnosperms, the ovules of flowering plants are enclosed within an ovary. After fertilization, the ovules become seeds, and the ovary develops into a fruit. 6 Briefly explain the life cycle of a flowering plant and describe double fertilization. ■ The sporophyte generation is dominant in flowering plants; gametophytes are extremely reduced in size and nutritionally dependent on the sporophyte generation. Flowering plants are heterosporous and produce microspores and megaspores within the flower. ■ Each microspore develops into a pollen grain (immature male gametophyte). One of each four megaspores produced by meiosis ■

■

develops into an embryo sac (female gametophyte). Within the embryo sac, the egg cell and the central cell with two polar nuclei participate in fertilization. Double fertilization, which results in the formation of a diploid zygote and triploid endosperm, is characteristic of flowering plants. Explore plant life cycles by clicking on the figures in CengageNOW.

7 Contrast monocots and eudicots, the two largest classes of flowering plants. ■ Most monocots (class Monocotyledones) have floral parts in threes, and their seeds each contain one cotyledon. The nutritive tissue in their mature seeds is endosperm. ■ Eudicots (class Eudicotyledones) usually have floral parts in fours or fives or multiples thereof, and their seeds each contain two cotyledons. The nutritive organs in their mature seeds are usually the cotyledons, which have absorbed the nutrients in the endosperm. 8 Discuss the evolutionary adaptations of flowering plants. ■ Flowering plants reproduce sexually by forming flowers. After double fertilization, seeds form within fruits. Flowering plants have efficient water-conducting vessel elements in their xylem and efficient carbohydrate-conducting sieve tube elements in their phloem. Wind, water, insects, or other animals transfer pollen grains in various flowering plants. Learn more about floral structure by clicking on the figure in CengageNOW.

28.4 (page 595) 9 Summarize the evolution of gymnosperms from seedless vascular plants and trace the evolution of flowering plants from gymnosperms. ■ Seed plants arose from seedless vascular plants. Progymnosperms were seedless vascular plants that had megaphylls and “modern” woody tissue. Progymnosperms probably gave rise to conifers as well as to seed ferns, which in turn likely gave rise to cycads and ginkgo. ■ The evolution of the gnetophytes is unclear, although molecular data indicate that they are closely related to conifers. ■ Flowering plants probably descended from ancient gymnosperms that had specialized features, such as leaves with broad, expanded blades and closed carpels. Flowering plants likely arose only once— that is, there is only one line of evolution from the gymnosperms to the flowering plants.

T E S T YO U R U N D E R S TA N D I N G 1. Seed plants lack which of the following structure(s)? (a) ovules surrounded by integuments (b) microspores and megaspores (c) vascular tissues (d) a large, nutritionally independent sporophyte (e) a large, nutritionally independent gametophyte 2. Conifers, cycads, ginkgo, and gnetophytes are collectively called (a) club mosses (b) gymnosperms (c) angiosperms (d) eudicots (e) seedless vascular plants 3. The immature male gametophytes of pine are called (a) ovules (b) stamens (c) seed cones (d) pollen grains (e) polar nuclei 4. The transfer of pollen grains from the male to the female reproductive structure is known as (a) pollination (b) fertilization

(c) embryo sac development (d) seed development (e) fruit development 5. Motile sperm cells are found as vestiges in these two gymnosperm groups: (a) monocots, eudicots (b) gnetophytes, conifers (c) gnetophytes, flowering plants (d) cycads, conifers (e) cycads, ginkgo 6. There are at least _____ species of flowering plants worldwide. (a) 235 (b) 3000 (c) 30,000 (d) 300,000 (e) 3,000,000 7. A simple pistil consists of a single (a) calyx (b) carpel (c) ovule (d) filament (e) petal

8. A flower that lacks stamens is both _____ and _____. (a) complete; imperfect (b) incomplete; perfect (c) complete; perfect (d) incomplete; imperfect

11. This flowering plant may be the nearest living relative to the ancestor of all flowering plants. (a) Amborella (b) Archaeopteris (c) Gnetum (d) water lily (e) Archaeanthus

9. After fertilization, the _____ develop(s) into a fruit and the _____ develop(s) into a seed. (a) ovary; ovule (b) polar nuclei; ovule (c) ovary; endosperm (d) ovule; ovary (e) ovule; polar nuclei

12.

10. The female gametophyte in flowering plants is also called the (a) polar nuclei (b) anther (c) embryo sac (d) endosperm (e) sporophyll

This cross section through an ovary reveals that the pistil is (a) simple, with one carpel (b) compound, with two fused carpels (c) compound, with three fused carpels (d) compound, with six separate carpels (e) compound, with six fused carpels

CRITICAL THINKING 1. How are cones and flowers alike? How are they different? (Hint: Your answer should consider microspores/megaspores and seeds.) 2. How do the life cycles of seedless plants (see Chapter 27) and seed plants differ? In what fundamental way are they alike?

6. You are given a plant that you have never seen before (see figure). Is it a gymnosperm or angiosperm? A monocot or eudicot? What are the features that helped you make these determinations?

4. EVOLUTION LINK. Contrast the algae, mosses, ferns, gymnosperms, and angiosperms with respect to their dependence on water as a transport medium for reproductive cells. Suggest a hypothesis to explain how the differences might be adaptive to living on land. 5. EVOLUTION LINK. Where would you place the progymnosperms on Figure 28-2? Explain your reasoning.

Carlyn Iverson

3. EVOLUTION LINK. Most flowers contain both male and female reproductive structures, in contrast to the cones of gymnosperms, which are either male or female. Explain how bisexual flowers might be an advantageous evolutionary adaptation to the flowering plants that possess them.

Additional questions are available in CengageNOW at www.cengage.com/ login.

29

The Fungi

Jeff Lepore/Photo Researchers, Inc.

Fungal spores. The rounded earthstar (Geastrum saccatum) releases a puff of microscopic spores after the sac, which is about 1.3 cm (0.5 in) wide, is hit by a raindrop. This fungus is common in leaf litter under trees throughout North America.

KEY CONCEPTS 29.1 Fungi are eukaryotic heterotrophs that absorb nutrients from their surroundings; most grow as multicellular filaments called hyphae that form a tangled mass called a mycelium. 29.2 Most fungi have complex life cycles, and reproduce both asexually and sexually by means of spores.

29.3 According to current hypotheses, fungi evolved from

M

ushrooms, morels, and truffles, delights of the gourmet, share a recent common ancestry with baker’s yeast, the black mold that forms

on stale bread, and the mildew that collects on damp shower curtains. All of these life-forms are classified as fungi, a diverse group of about 100,000 described species, most of which are terrestrial. Some mycologists, biologists who study fungi, estimate that there may be as many as 1.5 million species of fungi. Biologists who use the kingdom as a taxon group these organisms in kingdom Fungi, one of the eukaryotic kingdoms. Fungi grow best in moist habitats, but they are found universally

a unicellular, flagellate protist and diverged into several groups.

wherever organic material is available. They require moisture to grow,

29.4 Fungi are of major ecological importance; they are

and they can obtain water from a humid atmosphere as well as from the

decomposers and they form symbiotic relationships with other organisms.

medium on which they live. When the environment becomes dry, fungi

29.5 Fungi are of major economic, biological, and medical importance.

survive by going into a resting stage or by producing spores (see photograph) that resist desiccation (drying out). Some fungi grow to enormous size. In Washington State, a fungal clone (Armillaria ostoyae) has been identified that covers more than 1500 acres. This giant fungus, which is mainly underground, developed from a single spore that germinated more than a thousand years ago. The fungus has fragmented and is no longer one continuous body.

Like prokaryotes, most fungi are decomposers that obtain nutrients and energy from dead organic matter. They are vital members of ecosystems because they break down the organic compounds found in dead organisms, leaves, garbage, sewage, and other waste. When they decompose organic compounds, carbon and other elements are released into the environment, where they are recycled. Many fungi form vital symbiotic associations. For example, most terrestrial plants have fungal partners that live in close association with their roots. The fungi help the plants obtain phosphate ions and other needed minerals from the soil. In exchange, the plants provide the fungi with organic nutrients. Some fungi live symbiotically with algae and cyanobacteria as lichens. Oth-

Fungi absorb food from the environment Like animals, fungi are heterotrophs. For their nutritional and energy needs, they depend on preformed carbon molecules produced by other organisms. However, fungi do not ingest food and then digest it in the body as animals do. Instead, they infiltrate a food source and secrete digestive enzymes onto it. Digestion takes place outside the body. When complex molecules are broken down into smaller compounds, the fungus absorbs the predigested food into its body. The fungus is very efficient at absorbing nutrients and growing. It rapidly converts nutrients into new cell material. If excessive amounts of nutrients are available, fungi store them, usually as lipid droplets or glycogen.

ers are parasites and pathogens that cause disease in animals

Fungi have cell walls that contain chitin

or plants.

29.1 CHARACTERISTICS OF FUNGI ■ ■ LEARNING OBJECTIVES 1 2

Describe the distinguishing characteristics of fungi. Describe the body plan of a fungus.

All fungi (sing., fungus) are eukaryotes; their cells contain membraneenclosed nuclei, mitochondria, and other membranous organelles. Although they vary strikingly in size and shape, fungi share certain key characters, including their way of obtaining nutrition. The optimum pH for most fungal species is about 5.6, but various fungi can tolerate and grow in environments where the pH ranges from 2 to 9. Many fungi are less sensitive to high osmotic pressures than are bacteria. As a result, they can grow in concentrated salt solutions or in sugar solutions such as jelly, which discourage or prevent bacterial growth. Fungi also thrive over a wide temperature range. Even refrigerated food may be invaded by fungi.

Like the cells of bacteria, certain protists, and plants, fungal cells are enclosed by cell walls during at least some stage in their life cycle. Fungal cell walls, however, have a different chemical composition from cell walls of other organisms. In most fungi, the cell wall consists of complex carbohydrates, including chitin, a polymer that consists of subunits of a nitrogen-containing sugar (see Fig. 3-11). Chitin is also a component of the external skeletons of insects and other arthropods. Chitin is resistant to breakdown by most microorganisms.

Most fungi consist of a network of filaments The simplest fungi are the yeasts, which are unicellular, with a round or oval shape. Yeasts are widely distributed in the soil; on leaves, fruits, and cured meats; and on and in our bodies. The importance of some yeasts in medicine, biological research, and in the food industry is discussed later in this chapter. Most fungi are multicellular. The body consists of long, branched, threadlike filaments called hyphae (sing., hypha) (FIGS. 29-1a and b).

(c) A hypha divided into cells by septa. Each cell is monokaryotic (has one nucleus). In some taxa the septa are perforated, as shown.

G. T. Cole / University of Texas / Biological Photo Service

© Sidney Moulds / Photo Researchers, Inc.

Hyphae

25 μm

(a) A mycelium. A mass of hyphae has formed a mycelium (white area). Fruiting bodies, called sporangia, are visible as gray areas.

(b) SEM of a mycelium. The fungus Blumeria graminis growing on a leaf (darker area underneath the mycelium ).

(d) A septate hypha. In this hypha, each cell is dikaryotic (has two nuclei). (e) A coenocytic hypha.

FIGURE 29-1 Fungus body plan Spores carried by the air settle on food. They germinate and produce a mass of threadlike filaments called hyphae. The hyphae penetrate the food to obtain nourishment, and produce a mass of hyphae called the mycelium. Eventually, specialized hyphae grow from the mycelium giving rise to erect stalks with fruiting bodies (sporangia) at their tops. When the walls of the sporangia rupture, new spores are released into the air.

Hyphae consist of tubular cell walls surrounding the plasma membranes of the fungal cells. Hyphae are an adaptation to the fungal mode of nutrition. Growth occurs at the tips of the hyphae; as the hyphae elongate, the fungus grows into and infiltrates food sources. The fungus absorbs nutrients through its very large surface area. As hyphae grow, they form a tangled mass or tissuelike network, called a mycelium (pl., mycelia). Fungi that form mycelia are called molds. The cobweb-like mold sometimes seen on bread is the mycelium of a fungus. What is not seen is the extensive mycelium that grows into the bread. Depending on environmental conditions, some fungi can alternate between a yeast phase and a phase in which they produce hyphae. In most fungi, hyphae are divided by cross walls, called septa (sing., septum), into individual cells containing one or more nuclei (FIGS. 29-1c and d). As we will discuss, the presence of septa is an important character in the two largest fungal phyla (which include the most complex fungi). The septa of many fungi are perforated by a pore that may be large enough to permit organelles to flow from cell to cell. Some fungi, called coenocytic fungi, lack septa. In these species, nuclear division is not followed by cytoplasmic division. As a result, a coenocytic fungus is one elongated, multinucleated, giant cell (FIG. 29-1e).

Fungi produce spores either sexually or asexually. With asexual reproduction, new individuals are produced quickly, but there is little genetic variability. Sexual reproduction involves meiosis and generates new genotypes. Spores are usually produced on specialized aerial hyphae or in fruiting structures. When positioned up above the ground, spores can be easily dispersed. Structures in which spores are produced are called sporangia (sing., sporangium). The aerial hyphae of some fungi produce spores in large, complex reproductive structures, referred to as fruiting bodies. The familiar part of a mushroom is a large fruiting body. We do not normally see the bulk of the fungus, a nearly invisible mycelium buried out of sight in the rotting material or soil on which it grows.

Many fungi reproduce asexually Yeasts reproduce asexually, primarily by forming buds that pinch off from the parent cell (FIG. 29-3). Many species of multicellular fungi also reproduce asexually. Spores are produced by

Review ■ ■

What characters distinguish fungi from other organisms? How does the body of a yeast differ from that of a mold?

Bud development

29.2 FUNGAL REPRODUCTION ■ ■ LEARNING OBJECTIVE 3

(a) Yeasts can reproduce asexually by budding.

Describe the life cycle of a typical fungus, including sexual and asexual reproduction.

Most fungi reproduce by means of microscopic spores, reproductive cells that can develop into new organisms. In most groups, spores are nonmotile. They are dispersed by wind, water, or animals. The air we breathe is filled with hundreds of thousands of fungal spores. When a spore germinates, it gives rise to a hypha, which then develops into a mycelium (FIG. 29-2).

SciMat/ Photo Researchers, Inc.

Spore

5 μm

Hypha

Mycelium

FIGURE 29-2 Germination of a spore to form a mycelium

(b) SEM (color enhanced) of Saccharomyces cerevisiae. Also known as baker's or brewer's yeast, this fungus can ferment sugar, producing carbon dioxide and alcohol in the process. It is used in brewing beer, in wine production, and in baking bread. Note that several yeast cells are budding, a form of asexual reproduction.

FIGURE 29-3 Yeasts are unicellular fungi

mitosis and then released into the air or water. Conidiophores (from the Greek, meaning “dust-bearers”) are specialized hyphae that produce asexual spores called conidia (sing., conidium). The arrangement of conidia on conidiophores varies from species to species.

KEY POINT

Most fungi reproduce sexually Fungi are a diverse group with many variations in their life cycles. Most fungi (but not all) reproduce both asexually and sexually. FIGURE 29-4 illustrates a generalized life cycle. Many fungal species reproduce sexually when they come into contact with other mating

Most fungi can reproduce both asexually (which allows rapid proliferation) and sexually (which produces new genotypes).

7

Large numbers of haploid (n) spores are produced by mitosis.

8

Spore germinates and forms mycelium by mitosis.

2

Mycelia of two different mating types fuse at their tips, and plasmogamy (fusion of cytoplasm) occurs.

Asexual reproduction

1

6

Spores germinate and form mycelia by mitosis.

Mycelia

Spores are released. Spores

Sexual reproduction Haploid stage (n)

5

Plasmogamy

Dikaryotic stage (n + n)

Meiosis results in four genetically different haploid (n) nuclei. Spores develop around nuclei.

Diploid stage (2n) 3

Zygote nucleus (2n) Dikaryotic mycelium Meiosis

Karyogamy

4

FIGURE 29-4 The basic sequence of events in most fungal life cycles

Karyogamy (fusion of nuclei) occurs (black arrow), forming a diploid (2n) zygote nucleus.

Dikaryotic (n + n) mycelium develops.

types. In contrast to the majority of animal and plant cells, most fungal cells contain haploid nuclei. In sexual reproduction, the hyphae of two genetically compatible mating types come together, and their cytoplasm fuses, a process called plasmogamy. The resulting cell has two haploid nuclei, one from each fungus. This cell gives rise by mitosis to other cells with two nuclei. At some point the two haploid nuclei fuse. This process, called karyogamy, results in a cell containing a diploid nucleus known as a zygote nucleus. In some groups, the zygote nucleus is the only diploid nucleus. In the two largest fungal phyla, the ascomycetes and basidiomycetes (discussed later in this chapter), plasmogamy occurs (hyphae fuse), but karyogamy (fusion of the two different nuclei) does not follow immediately. For a time, the nuclei remain separate within the fungal cytoplasm. Hyphae that contain two genetically distinct, sexually compatible nuclei within each cell are described as dikaryotic (see Fig. 29-1d). This condition is referred to as n + n rather than 2n because there are two separate haploid nuclei. Hyphae that contain only one nucleus per cell are described as monokaryotic. The presence of a dikaryotic stage is an important defining character of the ascomycetes and basidiomycetes. Fungi communicate chemically by secreting signaling molecules called pheromones. At least one pheromone has been identified in each major fungal group. The pheromone binds with a compatible receptor on a different mating type. For example, in the zygomycetes, a pheromone induces the formation of specialized aerial hyphae. Another pheromone causes the tips of aerial hyphae of opposite mating types to grow toward each other and fuse prior to sexual reproduction.

Review ■ ■

How is a monokaryotic nucleus different from a dikaryotic nucleus? Draw a generalized life cycle of a fungus.

29.3 FUNGAL DIVERSITY ■ ■ LEARNING OBJECTIVES 4 Give arguments to support the hypothesis that fungi are opisthokonts, 5

more closely related to animals than to plants. Give arguments to support the hypothesis that chytrids may have been the earliest fungal group to evolve from the most recent common ancestor of fungi.

6 List distinguishing characteristics, describe a typical life cycle, and give examples of each of the following fungal groups: chytridiomycetes, zygomycetes, glomeromycetes, ascomycetes, and basidiomycetes.

For centuries, biologists classified fungi in the plant kingdom. Like plants, fungi have cell walls and vacuoles and are sessile, that is they cannot move around from place to place. Also, like plants, many types of fungi inhabit the soil. However, systematists began to question this classification, and in 1969, R. H. Whittaker proposed that fungi be assigned to a separate kingdom—Fungi. Unlike plants, fungal cell walls do not contain cellulose. Rather, they contain chitin, a polysaccharide found in insect skeletons. The fungal mode of nutrition is also very different from that of plants. Unlike plants, fungi cannot produce their own organic materials

from a simple carbon source (carbon dioxide). Like animals, fungi are heterotrophs. As with systematics for other kingdoms, fungal systematics is a challenging and continuously changing process. For example, slime molds and water molds were formerly classified as fungi but are now assigned to the protist kingdom (see Chapter 26).

Fungi are assigned to the opisthokont clade Systematists hypothesize that the common ancestor of all plants, fungi, and animals was a flagellated ancient protist. As discussed in Chapter 26, amoebozoa (a group of amoebas), fungi, animals, and a few protists, including the choanoflagellates (a group of flagellate protists), form a monophyletic “supergroup,” the unikonts. Within this supergroup, the clade opisthokonts includes the choanoflagellates, the animals, and the fungi. Both genetic and structural similarities support this grouping. Like animals, fungi have platelike cristae in their mitochondria. Another key character shared by members of this clade is that flagellate cells propel themselves with a single posterior flagellum. In other eukaryote groups, flagellate cells move by means of one or more anterior flagella. Based on structural characters and on molecular data, systematists now view fungi as more closely related to animals than to plants.

Diverse groups of fungi have evolved Fossil evidence has not been very helpful to systematists studying evolutionary relationships among fungal groups. Most fossilized fungi recovered to date have been microscopic. For example, fossilized fungal spores have been found in amber more than 225 million years old, and fossils of hyphae associated with cyanobacteria or algae have been dated as more than 550 million years old. Few large fungal fossils, such as mushrooms, have been found. Historically, fungi have been classified mainly on the characteristics of their sexual spores and fruiting bodies. More recently, molecular data, such as comparative DNA and RNA sequences, have helped clarify relationships among fungal groups. Currently, many mycologists assign fungi to five main groups: Chytridiomycota, Zygomycota, Glomeromycota, Ascomycota, and Basidiomycota (FIG. 29-5 and TABLE 29-1). Some biologists consider each of these groups to be a phylum. However, some of these groups are not monophyletic (see Chapter 23), and mycologists are in the process of assigning fungi to additional clades. In this edition of Biology, we will discuss fungi in the five phyla shown in Figure 29-5 and listed in Table 29-1. Microsporidia, a group of intracellular parasites, are classified in this text with the zygomycetes, although in the future they may be assigned to their own taxon. About 95% of all named fungi have been assigned to phyla Ascomycota and Basidiomycota. These phyla are considered sister taxa because they share a more recent common ancestor with each other than either does with any other group. Fungi of phyla Ascomycota and Basidiomycota have septate hyphae and a dikaryotic stage during the sexual part of their life cycle. Until recently, mycologists had assigned about 25,000 species of fungi that did not fit into the major groups to a group called deuteromycetes (phylum Deuteromycota). This was a polyphyletic group (members did not share a recent common ancestor) lumped

Basidiomycetes

Ascomycetes

Glomeromycetes

Zygomycetes

Chytrids

Evolution of ascospores Evolution of basidiospores Evolution of dikaryotic stage

Loss of flagellum Flagellate common ancestor

FIGURE 29-5 Cladogram of currently recognized groups of fungi This cladogram shows phylogenetic relationships among living fungi, based on comparisons of ribosomal and nuclear gene sequence data for many species. The chytrids were the lineage that branched off first during fungal evolution. Note that ascomycetes and basidiomycetes are sister clades. Remember that the phylogeny of fungi is a work in process.

together simply as a matter of convenience. Mycologists classified fungi as deuteromycetes if no sexual stage had been observed for them at any point during their life cycle. Some of these fungi have lost the ability to reproduce sexually, whereas others reproduce sexually only rarely. Most fungi classified as deuteromycetes reproduce only by means of conidia, as ascomycetes do. Mycolo-

TABLE 29-1

gists have identified relationships between deuteromycetes and their sexually reproducing relatives, based on DNA comparisons among various species. Most of the deuteromycetes have been reassigned to phylum Ascomycota, and a few reassigned to phylum Basidiomycota. (By convention, the asexual stage is still identified as a deuteromycete.)

Chytrids have flagellate spores At one time, biologists considered the chytrids, also known as chytridiomycetes (phylum Chytridiomycota), to be funguslike protists, similar in many respects to the water molds. However, both structural and molecular characters indicate that the approximately 1000 species of chytridiomycetes are members of kingdom Fungi. Like fungi, their cell walls contain chitin, and molecular comparisons, particularly of DNA and RNA sequences, have provided compelling evidence that chytrids are indeed fungi. However, recent comparisons of rRNA sequences suggest that chytrids are not a monophyletic group and they may be divided into four clades. Chytrids are small, relatively simple fungi that inhabit ponds and damp soil. A few species have been found in salt water. Most chytrids are decomposers that degrade organic matter. However, a few species cause disease in plants and animals. A parasitic chytrid has been partly responsible for declining amphibian populations. Infected frogs have been identified in many parts of the world (see Inquiring About: Declining Amphibian Populations, in Chapter 57). Most chytrids are unicellular or composed of a few cells that form a simple body, called a thallus. The term thallus describes the simple body plan of certain algae, fungi, and plants. The thallus may have slender extensions, called rhizoids, that anchor it to a food source and absorb food (FIG. 29-6). Chytrids are the only fungi that have flagellate cells. Their spores bear a single, posterior flagellum. Sexual reproduction has not been identified in most chytrids. Species that do reproduce sexually have flagellate gametes.

Characteristics of Currently Recognized Phyla of Fungi

Phylum and Common Types

Asexual Reproduction

Sexual Reproduction

Other Key Characters

Chytridiomycota (chytrids or chytridiomycetes) Allomyces

Flagellate, diploid zoospores produced by mitosis in zoosporangia

Flagellate, haploid gametes in some species

Haploid zoospores produced in resting sporangia; form haploid thallus

Zygomycota (zygomycetes) Black bread mold. Microsporidia are classified with the zygomycetes

Haploid spores produced in sporangia

Zygospores develop in zygosporangia

Important decomposers; some are insect parasites. Microsporidia are opportunistic pathogens that infect animals.

Glomeromycota (glomeromycetes)

Large, multinucleate blastospores

Has not been observed

Form arbuscular mycorrhizae with plant roots

Ascomycota (ascomycetes) Yeasts, powdery mildews, molds, morels, truffles

Conidia pinch off from conidiophores

Ascospores develop in asci

Have a dikaryotic stage; form important symbiotic relationships as lichens and mycorrhizae

Basidiomycota (basidiomycetes or club fungi) Mushrooms, bracket fungi, puffballs, rusts, smuts

Uncommon

Basidiospores develop on club-shaped basidia

Have a dikaryotic stage; many form mycorrhizae with tree roots

lutionary history, other fungal groups apparently lost the ability to produce motile cells, perhaps during the transition from aquatic to terrestrial habitats.

John Taylor/Visuals Unlimited, Inc.

Zygomycetes reproduce sexually by forming zygospores

5 μm

FIGURE 29-6 Chytrid Nomarski differential interference micrograph of a common chytrid (Chytridium convervae). Many chytrids have a microscopic body form consisting of a rounded, coenocytic thallus and branched rhizoids that superficially resemble roots. The rhizoids may anchor the chytrid thallus and absorb predigested food.

Some chytrids produce branched, coenocytic mycelia. Allomyces, a large, common chytrid, has an unusual life cycle compared to that of most fungi. It undergoes an alternation of generations (common in plants, but rare in fungi), spending part of its life as a multicellular haploid (n) thallus and part as a multicellular diploid (2n) thallus (FIG. 29-7). The haploid and diploid thalli are similar in appearance. At the tips of its branches, the haploid thallus bears two types of sporangia, structures in which gametes form by mitosis. Each sporangium produces a different type of flagellate gamete. Each type of gamete secretes a pheromone that attracts the other type. The two gametes fuse, and plasmogamy and karyogamy occur, resulting in a motile zygote. Each zygote can develop into a diploid thallus. The thallus bears two kinds of spore cases, zoosporangia and resting sporangia. Zoosporangia produce flagellate diploid zoospores that develop into new diploid thalli. Meiosis occurs within resting sporangia, producing haploid zoospores. Each zoospore has the potential to develop into a haploid thallus. Molecular evidence suggests that chytrids were probably the earliest fungal group to evolve. Chytrids produce flagellate cells at some stage in their life history, a character that systematists trace to the protist that was the common ancestor of all opisthokonts. No other fungal group has flagellate cells. At some point in their evo-

Mycologists have named more than 1100 species of zygomycetes (phylum Zygomycota). Of all the fungi, members of this taxon appear most closely related to the chytrids. However, zygomycetes are not a monophyletic group, and as mycologists learn more about fungal relationships, they may divide this phylum into several taxa or reassign its members to other existing phyla. Most zygomycetes are decomposers that live in the soil on decaying plant or animal matter (FIG. 29-8). Some zygomycetes form a type of symbiotic association (mycorrhizal relationship) with plant roots. (Recall that a symbiotic association is an intimate relationship between organisms of different species.) A few species cause disease in plants and animals, including humans. During sexual reproduction, zygomycetes produce sexual spores, called zygospores. The zygospores are typically produced in spore sacs called zygosporangia. The hyphae in zygomycetes are coenocytic; that is, they lack regularly spaced septa. However, septa do form to separate the hyphae from reproductive structures. Perhaps the most familiar zygomycete is the black bread mold, Rhizopus stolonifer, a decomposer that breaks down bread and other foods. If preservatives are not added, bread left at room temperature for a few days often becomes covered with a black, fuzzy growth. Bread becomes moldy when a spore falls on it and then germinates and grows into a mycelium (FIG. 29-9 on page 610). Hyphae penetrate the bread and absorb nutrients. Eventually, certain hyphae grow upward and develop sporangia at their tips. Clusters of more than 50,000 black asexual spores can develop within each sporangium. The spores are released when the delicate sporangium ruptures. The spores give the black bread mold its characteristic color. Sexual reproduction in the black bread mold occurs when the hyphae of two different mating types, designated plus (+) and minus (−), grow into contact with one another. The bread mold is heterothallic, meaning that an individual fungal hypha mates only with a hypha of a different mating type. That is, sexual reproduction occurs only between a member of a (+) strain and one of a (−) strain, not between members of two (+) strains or members of two (−) strains. Because there are no physical differences between the two mating types, it is not appropriate to refer to them as “male” and “female.” When hyphae of opposite mating types grow in close proximity, they signal one another with pheromones. In response to these chemical signals, the tips of the hyphae come together and form gametangia, which serve as gametes. Plasmogamy occurs as the gametangia fuse. Then karyogamy occurs as the (+) and (−) nuclei fuse to form the diploid zygote nucleus. The zygote develops into a zygospore. Zygospores are encased in a thick protective zygosporangium. The zygospore may lie dormant for several months. It can survive desiccation and extreme temperatures. Meiosis probably occurs at or just before germination of the zygospore.

Ascomycetes

Basidiomycetes

Chytrids have flagellate reproductive cells.

Glomeromycetes

Zygomycetes

Chytrids

KEY POINT

Mature haploid thallus Sporangium

Flagellate common ancestor

2 1

Haploid zoospore grows into haploid thallus. Haploid

Haploid thallus produces two types of gametes by mitosis. Gamete type A

zoospore

6

Gamete type B

Haploid zoospores are produced by meiosis. Resting sporangium

SEXUAL REPRODUCTION HAPLOID (n) GENERATION 3

DIPLOID (2n) GENERATION

Plasmogamy and karyogamy

Meiosis 5

Meiosis occurs in resting sporangia.

Motile zygote

Resting sporangium

Zoosporangium

ASEXUAL REPRODUCTION (by mitosis)

Diploid zoospore

FIGURE 29-7 Life cycle of the chytrid Allomyces arbuscula Allomyces alternates between haploid and diploid stages, which are similar in appearance.

Gametes fuse and their nuclei fuse, producing flagellate zygote.

4

Zygote germinates and develops into diploid thallus.

7

Zoosporangia produce flagellate diploid zoospores by mitosis. Zoospores give rise to new diploid thalli.

Cabisco/Visuals Unlimited

FIGURE 29-8 Pilobolus, a zygomycete that grows in animal dung Stalked sporangia of Pilobolus protruding from a pile of dung, which contains an extensive mycelium of the fungus. The stalked sporangia, which are 5 to 10 mm tall, act like shotguns and forcefully discharge sporangia (the black tips) away from the dung onto nearby grass. When animals such as cattle or horses eat the grass, the spores pass unharmed through the animal’s digestive tract and are deposited in a fresh pile of dung.

When the zygospore germinates, an aerial hypha develops with a sporangium at the tip. Mitosis within the sporangium produces haploid spores. These spores may be all (+) spores, all (−) spores, or a mixture of (+) and (−) spores. When released, the spores germinate to form new hyphae. Only the zygote and zygospore of a black bread mold are diploid; all the hyphae and the asexual spores are haploid.

Microsporidia have been a taxonomic mystery Microsporidia are small, unicellular parasites that infect eukaryotic cells. They are opportunistic pathogens that infect animals. For example, microsporidia infect people with compromised immune systems, such as those with AIDS. Microsporidia cause a variety of diseases involving many organ systems, and some species cause lethal infections. Some species of microsporidia appear to be host-specific. Microsporidia have two developmental stages inside their host: a feeding stage and a reproductive stage. Some microsporidian species divide into two cells by binary fission, and others divide into several cells. Some species undergo nuclear fusion and meiosis before producing spores. The spores, which have thick protective walls, can pass from cell to cell inside the host or can be excreted in urine or through the skin. The spores, the only stage with distinct characters, are used to identify groups. Each spore is equipped with a unique structure, a long, threadlike polar tube. When the spore enters the gut of a new host, it discharges its polar tube and penetrates the lining of the gut. Acting as a hypodermic needle, the polar tube injects the contents of the spore into the host cell (FIG. 29-10). Microbiologists estimate that there may be more than a million species of microsporidia, but only about 1500 species have been named. Microsporidia were originally classified with yeasts and

bacteria. In 1976, they were assigned to the protozoa. In the 1980s, biologists viewed microsporidia as the most primitive example of a eukaryote. They were the smallest and simplest known eukaryotes, and the genomes of some species are smaller than most bacterial genomes. Microsporidia lack mitochondria, flagella, and Golgi complexes. Their ribosomes resemble those of prokaryotes. In 1998, British biologist Thomas Cavalier-Smith reassigned microsporidia to kingdom Fungi. Molecular studies show that microsporidia have gene sequences that indicate they originally had mitochondria. Microbiologists now generally agree that these organisms have become simpler as they adapted to their parasitic way of life. Other molecular studies have provided additional evidence for their taxonomic relationship with fungi. Establishing that they are closely related to fungi is important in developing medications that will be effective in treating microsporidian infections. Recent genome studies suggest that microsporidia descended from a zygomycete ancestor. For now, we classify the microsporidia with the zygomycetes, but in the future they may be assigned to a separate taxon.

Glomeromycetes are symbionts with plant roots Glomeromycetes (phylum Glomeromycota) have coenocytic (no septa) hyphae. They reproduce asexually with large, multinucleate spores called blastospores. Sexual reproduction has not been documented. Only about 200 species of glomeromycetes have been described. Glomeromycetes were previously considered zygomycetes, but in 2002 taxonomists determined that they form a separate monophyletic group. This phylogeny is based on molecular data. Glomeromycetes are symbionts that form intracellular associations with the roots of most trees and herbaceous plants. These symbiotic associations between the hyphae of certain fungi and the roots of plants are called mycorrhizae (from Greek words meaning “fungus roots”) (FIG. 29-11). Glomeromycetes extend their hyphae through the cell walls of root cells but often may not penetrate the plasma membrane. As each hypha pushes forward, the plasma membrane of the root cell surrounds it. Thus, the hyphae can be thought of as fingers pushing into a glove formed by the plasma membrane. Because they penetrate the cell wall, these fungi are referred to as endomycorrhizal fungi. The most widespread endomycorrhizae are called arbuscular mycorrhizae because the hyphae inside the root cells form branched, tree-shaped structures known as arbuscules (see Fig. 29-11 and Fig. 36-11b). The arbuscules are the sites of nutrient exchange between the plant and the fungus. Arbuscular mycorrhizae live entirely underground. In mycorrhizal relationships, the roots supply the fungus with sugars, amino acids, and other organic substances. The mycorrhizal fungus decomposes organic material in the soil and also benefits the plant by extending the reach of its roots. The slender mycelia are far thinner than roots and can extend into narrow spaces, absorbing nutrients that the plant could not capture on its own. Thus, with the help of the mycorrhizal fungus, the plant can also take in more nutrient minerals such as phosphorus and nitrogen. What we have just described is a mutualistic symbiotic association: both partners benefit. Studies show that if a plant grows

Ascomycetes

Basidiomycetes

Like most fungi, most zygomycetes reproduce both asexually and sexually.

Glomeromycetes

Zygomycetes

Chytrids

KEY POINT

Spore germinates

In asexual reproduction, certain hyphae form sporangia in which clusters of black, asexual, haploid spores develop. When released, they give rise to new hyphae.

2a

ASEXUAL REPRODUCTION (by spores) Haploid (n)

Flagellate common ancestor

Spores germinate and produce haploid mycelia.

1

Sporangia

Spores

+ –

SEXUAL REPRODUCTION 2b

Sporangium containing haploid spores produced by mitosis

Germination of zygospore

7

Meiosis occurs and zygospore germinates; hypha develops sporangium at its tip. 6

HAPLOID (n) STAGE

DIPLOID (2n) STAGE

Hyphae of (+) and (−) mating types grow toward one another.

Gametangia

+ –

Mature zygospore within zygosporangium

Meiosis

Zygospore develops from zygote; it is encased by thick-walled, black zygosporangium.

When (+) and (−) hyphae meet, they form 4 Plasmogamy occurs gametangia. as gametangia fuse.

Plasmogamy Karyogamy

5

+ –

Karyogamy occurs with nuclei fusing to form diploid zygote.

FIGURE 29-9 Animated Life cycle of a zygomycete, the black bread mold (Rhizopus stolonifer)

3

Microsporidian cell

Cells of root cortex

Root epidermis

Polar tube

Soil

Vesicle

Root hair

Host cell

1

2 Spore ejects 3 Infective Spore of microsporidium its polar tube cytoplasm is has coiled and penetrates injected into polar tube. host cell. host cell.

Arbuscule

Spore

Cortex cell Hypha of fungus

FIGURE 29-10 Infection by a microsporidium FIGURE 29-11 Arbuscular mycorrhizae

in phosphate-deficient soil or if it has a limited root system, its growth is enhanced by having a fungal partner. However, a plant in a phosphate-rich soil with a well-developed root system may not need fungal partners. For these plants, the fungus may be a parasite. Much remains to be learned about mycorrhizal associations. Other, yet unknown, benefits of the symbiosis may further define the association as mutualistic. For example, some fungi release alkaloids that protect the plants from herbivores and pathogens. Plants also exchange nutrients with one another through fungi that connect them. Scientists have discovered mycorrhizal fungi within ancient plant fossils in rocks that are about 400 million years old. These findings suggest that when plants moved onto the land, their fungal partners moved with them. In fact, the fungal partners may have been critical for early vascular plants to colonize the land because the fungal hyphae may have provided plants with water and minerals before their own root systems evolved.

Ascomycetes reproduce sexually by forming ascospores Ascomycetes (phylum Ascomycota) comprise a large group of fungi consisting of more than 32,000 described species. The diverse ascomycetes include most yeasts; the powdery mildews; most of the blue-green, pink, and brown molds that cause food to spoil; decomposer cup fungi; and the edible morels and truffles. The ascomycetes, more than any other group of fungi, impact humans. As we will discuss in a later section, ascomycetes are used to flavor cheeses, to bake bread (yeast), and to ferment alcohol. Some are enjoyed as foods (morels and truffles). Ascomycetes are used to produce antibiotics. They have also served as valuable model organisms for biologists studying cellular processes, including protein synthesis. Many fungi in this group form mycorrhizae with tree roots, and about 40% join with green algae or cyanobacteria to form lichens. On the negative side, ascomycetes cause

This mycelium has grown into a plant root. Its hyphae branch between the cells of the root. Hyphae have penetrated through the cell walls of two root cells and have branched extensively to form arbuscules. The tip of one hypha between root cells has enlarged and serves as a vesicle that stores food. The tip of a hypha in the soil has enlarged, forming a spore. The spaces between the root cells have been magnified for clarity.

most fungal diseases of plants and animals, including humans. For example, ascomycetes cause serious plant diseases such as Dutch elm disease, ergot disease on rye, powdery mildew on fruits and ornamental plants, and chestnut blight. Ascomycetes are sometimes referred to as sac fungi because their sexual spores are produced in microscopic sacs called asci (sing., ascus). Their hyphae usually have septa, but these cross walls have pores so that cytoplasm is continuous from one cell compartment to another. In most ascomycetes, asexual reproduction involves production of spores called conidia, which form at the tips of certain specialized hyphae known as conidiophores (FIG. 29-12). Production of these spores is a means of rapidly propagating new mycelia when environmental conditions are favorable. Conidia occur in various shapes, sizes, and colors in different species. The color of the conidia produces the characteristic blue-green, pink, brown, or other tints of many of these molds. Some species of ascomycetes are heterothallic. (Recall that heterothallic means that an individual fungal hypha mates only with a hypha of a different mating type.) Others are homothallic, which means they are self-fertile and have the ability to mate with themselves. In both heterothallic and homothallic ascomycetes, sexual reproduction takes place after two gametangia come together and their cytoplasm mingles. Let us examine the life cycle of a typical ascomycete (FIG. 29-13). In our example, plasmogamy takes place as hyphae of two different mating types come together and fuse. Within this fused structure, pairs of haploid nuclei, one from each parent hypha, associate but

Conidia

separates from the parent cell (see Fig. 29-3). Each bud can grow into a new yeast cell. Yeasts reproduce sexually by forming ascospores. During sexual reproduction, two haploid yeasts fuse, forming a diploid zygote. The zygote undergoes meiosis, and the resulting haploid nuclei are incorporated into ascospores. These spores remain enclosed for a time within the original cell wall, which corresponds to an ascus.

Biophoto Associates/Photo Researchers, Inc.

Basidiomycetes reproduce sexually by forming basidiospores

10 μm

FIGURE 29-12 Conidia SEM of Penicillium conidiophores, which resemble paintbrushes. Note the conidia pinching off from the tips of the “brushes.” Conidia are asexual reproductive cells produced by ascomycetes and some basidiomycetes. Biologists use the arrangement of conidia on conidiophores to identify species of these fungi.

do not fuse. New hyphae, with dikaryotic cells, develop from the fused structure. The hyphae branch repeatedly until the hyphal tips reach the site where asci will be produced. As the many sac-shaped asci develop, each containing two dissimilar nuclei (one from each parent), they are surrounded by intertwining haploid (monokaryotic) hyphae. These hyphae help make a fruiting body known as an ascocarp (FIG. 29-14a). Karyogamy occurs in each ascus. The two nuclei fuse and form a diploid zygote nucleus. The zygote nucleus then undergoes meiosis to form four haploid nuclei with different genotypes. One mitotic division of each of the four nuclei usually follows, resulting in eight haploid nuclei. Each haploid nucleus becomes incorporated into a thick-walled ascospore; thus, there are typically eight haploid ascospores within the ascus (FIG. 29-14b). The ascospores are usually released through a pore, slit, or hinged lid at the tip of the ascus. Air currents carry individual ascospores, often for long distances. If one lands in a suitable location, it germinates and forms a new mycelium. The fungus can reproduce asexually by producing conidia that can develop into new mycelia. Phylum Ascomycota includes more than 300 species of unicellular yeasts. Asexual reproduction of yeasts is mainly by budding; in this process a small protuberance (bud) grows and eventually

The more than 30,000 species of basidiomycetes (phylum Basidiomycota) include the largest and most familiar of the fungi: the mushrooms, bracket fungi, and puffballs (FIG. 29-15). Many basidiomycetes are decomposers that obtain nutrients by breaking down organic matter. Some species cause great economic loss because they cause dry rot in buildings. Certain basidiomycetes form mycorrhizae. Others, such as wheat rust and corn smut, infect important crops. A few basidiomycetes cause human disease. Sometimes called club fungi, basidiomycetes derive their name from their microscopic club-shaped basidia (sing., basidium). Basidia are comparable in function to the asci of ascomycetes. Each basidium is an enlarged hyphal cell that undergoes meiosis to form four basidiospores (FIG. 29-16). Note that basidiospores develop on the outside of a basidium, whereas ascospores develop within an ascus. Each individual fungus produces millions of basidiospores, and each basidiospore has the potential to give rise to a new primary mycelium. Hyphae of a primary mycelium consist of monokaryotic cells. The mycelium of a basidiomycete, such as the commonly cultivated mushroom Agaricus brunnescens, consists of a mass of white, branching, threadlike hyphae that live mostly underground. Septa divide the hyphae into cells, but as in ascomycetes, the septa are perforated and allow cytoplasmic streaming between cells. Let us examine the life cycle of a typical basidiomycete. Asexual reproduction is less common in basidiomycetes than in other groups, so we will focus here on sexual reproduction. We begin with two compatible primary mycelia (FIG. 29-17 on page 616). When in the course of its growth a hypha of a primary mycelium encounters a compatible monokaryotic hypha, typically of a different mating type, the two hyphae fuse (plasmogamy). As in the ascomycetes, the two haploid nuclei remain separate within each cell. In this way a secondary mycelium with dikaryotic hyphae is produced, in which each cell contains two haploid nuclei. The n + n hyphae of the secondary mycelium grow rapidly and extensively. When environmental conditions are favorable, the hyphae form compact masses, called buttons, along the mycelium. Each button grows into a fruiting body that we know as a mushroom. A mushroom is more formally referred to as a basidiocarp. Each basidiocarp consists of intertwined, matted hyphae, and has a stalk and a cap. The lower surface of the cap usually consists of many thin, perpendicular plates called gills that radiate from the stalk to the edge of the cap. Karyogamy takes place within the young basidia on the gills of the mushroom. The haploid nuclei fuse in the dikaryotic cells,

Basidiomycetes

Ascomycetes

Glomeromycetes

Zygomycetes

Ascomycetes produce asexual spores called conidia and sexual spores called ascospores.

Chytrids

KEY POINT

Conidia 10

In asexual reproduction, hyphae produce haploid conidia that can develop into new mycelia.

Germinating conidium

Flagellate common ancestor

ASEXUAL REPRODUCTION (by conidia) Conidiophore 9

8

When (+) mating type released, ascospores germinate and form new haploid mycelia.

Haploid (n)

1

(–) mating type

Plasmogamy

Each nucleus becomes incorporated into an ascospore.

2

SEXUAL REPRODUCTION HAPLOID (n) STAGE

Plasmogamy occurs as hyphae of the two mating types fuse and nuclei are exchanged.

DIKARYOTIC STAGE (n + n)

Mitosis produces eight haploid nuclei. Second meiotic division First meiotic division

Nuclei migrate

3

Mature ascus has eight haploid ascospores 7

Haploid mycelia of opposite mating types both produce coenocytic sexual hyphae.

DIPLOID (2n) Developing STAGE ascus with Nuclei fuse n + n nuclei

4

Hyphae form an ascocarp.

Zygote 6

Meiosis occurs, forming four haploid nuclei.

Meiosis

Ascocarp

Karyogamy

5

Karyogamy occurs in each ascus. Two haploid nuclei fuse, forming a diploid zygote nucleus.

Mycelium

FIGURE 29-13 Life cycle of a typical heterothallic ascomycete Sexual reproduction requires haploid mycelia of different mating types. Note the dikaryotic stage and the separation of plasmogamy and karyogamy. Steps 5 through 8 take place within an ascus in the ascocarp.

Dikaryotic hyphae form and produce asci.

© Robert and Linda Mitchell

© Ed Reschke

(a) The ascocarp (fruiting body) of the common brown cup (Peziza badio-confusa). This ascocarp is shaped like a saucer or bowl and is 3 to 10 cm (1 to 4 in) wide. It is found on damp soil in woods throughout North America. Photographed in Muskegon, Michigan.

100 μm

(b) Asci. Each ascus contains eight ascospores. Asci line the inner portion of the ascocarp.

FIGURE 29-14 Animated Sexual reproduction in the ascomycetes

forming diploid zygote nuclei. These are the only diploid cells that form during a basidiomycete’s life cycle. Meiosis then takes place, forming four haploid nuclei with different genotypes. These nuclei move to the outer edge of the basidium. Fingerlike extensions of the basidium develop, into which the nuclei and some cytoplasm move; each of these extensions becomes a basidiospore. A septum forms that separates the basidiospore from the rest of the basidium by a delicate stalk that breaks when the basidiospore is forcibly discharged. Each basidiospore can germinate and give rise to a primary mycelium.

Many basidiomycetes produce “fairy rings” in lawns and forests (FIG. 29-18 on page 617). A fairy ring may first appear as a dark green ring surrounding an inner brown circle. The size of the ring ranges from a few centimeters to more than 15 m (about 51 ft) in diameter. The green ring consists of grass, well nourished by the nutrients released as the fungi decompose organic material. Grass dies, producing the inner brown circle, because the mass of mycelia decreases the movement of water into the area. As the fungi grow outward, the circle widens. The rings grow a few centimeters to more than a meter per year. After rainfall or irrigation, a

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Cengage

FIGURE 29-15 Basidiomycete fruiting bodies

(a) Basidia line the gills of the Jack-o’-lantern mushroom (Omphalotus olearius ). The gills of this poisonous species produce a greenish glow in the dark. Each cap is about 15 cm (6 in) wide. Photographed at the base of an oak tree in Maryland, the Jack-o’lantern occurs throughout eastern North America and California.

ring of mushrooms may appear just outside the green circle. The name “fairy ring” comes from a legend that a ring of mushrooms appeared where fairies had danced in a circle the night before. Basidiospore

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What evidence supports the hypothesis that chytrids were the earliest fungal group to evolve from the common ancestor of fungi? What are the distinguishing characteristics of each of the following fungal groups: (1) zygomycetes, (2) glomeromycetes, (3) ascomycetes, and (4) basidiomycetes? How does the life cycle of a typical basidiomycete differ from that of a typical ascomycete? (Draw diagrams to support your answer.) Distinguish among (1) ascocarp, ascus, and ascospore and among (2) basidiocarp, basidium, and basidiospore.

29.4 ECOLOGICAL IMPORTANCE OF FUNGI ■ ■ LEARNING OBJECTIVES

5 μm

FIGURE 29-16 SEM of a basidium

7 Summarize the ecological significance of fungi as decomposers. 8 Describe the important ecological role of mycorrhizae. 9 Characterize the unique nature of a lichen.

Each basidium produces four basidiospores.

fungi degrade wastes and dead organisms, they release water, carbon (as CO2), and mineral components of organic compounds, and these elements are recycled (see biogeochemical cycles in Chapter 55). Without this continuous decomposition, essential nutrients would remain locked up in huge mounds of animal carcasses, feces, branches, logs, and leaves. These nutrients would be unavailable for use by new generations of organisms, and life would eventually cease. Fungi form important symbiotic relationships with animals, plants, bacteria, and protists. A three-way symbiotic relationship involving a fungus, a grass, and a virus has recently been reported. In the geothermal hot spots of Yellowstone National Park, a fungus infects the roots of the host grass. When infected with a specific virus, the fungus is heat tolerant and confers heat tolerance to the grass. When not infected by the virus, the fungus does not confer heat tolerance. The symbiotic relationships of fungi with other organisms have major effects on ecosystems. J. L. Lepore/Photo Researchers, Inc.

Richard D. Poe/Visuals Unlimited

Fungi make vital contributions to the ecological balance of our planet. Like bacteria, most fungi are free-living decomposers, chemoheterotrophs that absorb nutrients from organic wastes and dead organisms. For example, many fungal decomposers degrade cellulose and lignin, the main components of plant cell walls. When

(b) The elegant stinkhorn (Phallus ravenelii ) has a foul smell that attracts flies. The flies help disperse the slimy mass of basidiospores. Fruiting bodies of elegant stinkhorns grow to 18 cm (7 in) tall. Photographed in Pennsylvania.

Basidium

Biophoto Associates/Photo Researchers, Inc.

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(c) Turkey-tail (Trametes versicolor ) is a common bracket fungus. Bracket fungi grow on both dead and living trees and produce shelflike fruiting bodies. Basidiospores are produced in pores located underneath each shelf.

Fungi form symbiotic relationships with some animals Because animals do not have the enzymes necessary to digest cellulose and lignin, cattle and other grazing animals cannot, by themselves, obtain needed nutrients from the plant material they eat. Their survival depends on fungi that

Ascomycetes

Basidiomycetes

Basidiomycetes produce sexual basidiospores on the gills of basidiocarps (fruiting bodies).

Glomeromycetes

Zygomycetes

Chytrids

KEY POINT

1

Flagellate common ancestor

Basidiospores germinate and form primary mycelia.

2

Basidiospores released

Plasmogamy of primary mycelia occurs with the fusion of two (n) hyphae of different mating types.

Basidiospores forming

7

Plasmogamy

3

HAPLOID (n) STAGE Second meiotic division

First meiotic division

6

Meoisis occurs, producing four haploid nuclei that become basidiospores.

Meiosis

Fast-growing secondary mycelium is produced, composed of dikaryotic (n + n) hyphae.

DIKARYOTIC STAGE (n + n)

4

DIPLOID (2n) STAGE

Zygote Karyogamy

5

Gills

Basidia form along gills of basidiocarps. In each basidium karyogamy occurs, producing a zygote nucleus.

FIGURE 29-17 Animated Life cycle of a typical basidiomycete Note the dikaryotic stage and the separation of plasmogamy and karyogamy. Steps 5 and 6 take place within the basidia of the basidiocarp. Asexual reproduction is uncommon in this group.

Basidiocarps periodically develop from secondary mycelium.

Basidiocarp

Secondary mycelium

Wally Eberhart/Visuals Unlimited, Inc.

FIGURE 29-18 A fairy ring

inhabit their guts because fungi, like many other microorganisms, do have the enzymes that break down these organic compounds. The fungi benefit by living in a nutrient-rich environment. Fungi also form symbiotic associations with ants and termites. More than 200 species of ants farm fungi. Leaf-cutting ants bring leaves to their fungi and protect them from competitors and predators. The ants also disperse the fungi to new locations. In exchange, the fungi digest the leaves, providing nutrients for the ants. This symbiosis can involve other organisms. The farmed fungi can be infested by fungal parasites. In response, the ants culture bacteria (actinomycetes) that produce antibiotics to control these parasites. These symbiotic relationships, the most complex known, are the product of 50 million years of coevolution.

Mycorrhizae are symbiotic associations between fungi and plant roots Mycorrhizae occur in about 80% of plants (and more than 90% of all plant families). As discussed in the section on glomeromycetes, mycorrhizal fungi decompose organic material in the soil and increase the surface area of a plant’s roots so that the plant can absorb more water and mineral nutrients. In exchange, the roots supply the fungus with organic nutrients. In order to establish and maintain a symbiotic relationship, cells of the fungi and cells of the plant roots must communicate. For example, signaling molecules from the plant root cells stimulate fungal cells to shift to a presymbiotic growth phase in which their energy metabolism increases and their hyphae branch. The fungal cells then signal the root cells, activating a signaling pathway that activates gene expression in the root cells. The importance of mycorrhizae first became evident when horticulturalists observed that orchids do not grow unless an appropriate fungus lives with them. Similarly, many forest trees, such as pines, decline and eventually die from mineral deficiencies when transplanted to mineral-rich grassland soils that lack the appropriate mycorrhizal fungi. When forest soil containing the appropriate fungi or their spores is added to the soil around these trees, they quickly resume normal growth. Studies performed with various

types of plants, including cedar, have confirmed the role of mycorrhizae in plant growth (FIG. 29-19). As we have discussed, glomeromycetes form endomycorrhizal connections; they infiltrate the cells of plant roots. At least 5000 species of ascomycetes and basidiomycetes also form mycorrhizal connections, but their hyphae coat the plant root rather than penetrate its cells. They are referred to as ectomycorrhizae. Interestingly, researchers have shown that some mycorrhizal fungi harbor bacteria in their cytoplasm. Although the role of the bacteria is not yet clear, their presence suggests that they may be members of a three-way partnership: fungus, plant, and bacteria. Mycorrhizal fungi connect plants, allowing nutrient transfer among them. Scientists have measured the movement of organic materials from one tree species to another through shared mycorrhizal connections. Mycorrhizal fungi also release chemicals that protect the plant against herbivores and pathogens. Mycorrhizae improve the soil by decreasing water loss and erosion. Ecologists are studying the role of mycorrhizal fungi in reclaiming soils damaged by pollution. For example, mycorrhizae can modify toxic heavy metals, such as cadmium, so that plants cannot absorb them.

A lichen consists of two components: a fungus and a photoautotroph Although a lichen looks like a single organism, it is actually a dual organism, a combination of a fungus and a photoautotroph (FIG. 29-20a). Almost one-fifth of all known fungal species form these symbiotic relationships. About 14,000 kinds of lichens have been described. Fossils suggest that fungi developed symbiotic partnerships with photoautotrophs before the evolution of vascular plants. The photoautotrophic component of a lichen is either a green alga, a cyanobacterium, or both. The fungus is most often an ascomycete, although a basidiomycete is the fungal partner in some tropical lichens. Most photoautotrophic organisms found in lichens also occur as free-living species in nature, but the fungal components are generally found only as a part of the lichen. Typically, the fungus forms most of the lichen thallus (body). The fungus surrounds hundreds of photosynthetic partners and holds them in place. Lichens are named for the fungal component. In the laboratory, researchers can isolate the fungal and photoautotrophic components of some lichens and grow them separately in appropriate culture media. The photoautotroph grows more rapidly when separated, whereas the fungus grows more slowly and requires many complex carbohydrates. Neither organism resembles a lichen in appearance when grown separately. The photoautotroph and fungus can be reassembled as a lichen thallus, but only if they are placed in a culture medium under conditions that cannot support either of them independently. What is the nature of this partnership? The lichen was originally considered a definitive example of mutualism. The photoautotroph carries on photosynthesis, producing energy-rich carbon compounds for both members of the lichen. It is unclear how the photoautotroph benefits from the relationship. Some biologists have suggested that the photoautotroph obtains water and nutrient minerals from the fungus as well as protection against desiccation.

KEY EXPERIMENT QUESTION: Is plant growth affected by fungi in the soil? HYPOTHESIS: In low phosphorus soils, plants that form mycorrhizal associations with fungi exhibit enhanced growth.

Courtesy of Randy Molina, U.S. Forest Service

Courtesy of Randy Molina, U.S. Forest Service

EXPERIMENT: Western red cedar seedlings (Thuja plicata) were selected for this study. Control seedlings were grown in low phosphorus soil in the absence of the fungus. Experimental seedlings of the same age as the control plants were grown under the same conditions as the controls, except that their roots formed mycorrhizal associations.

(a) No mycorrhizal associations. Control seedlings were grown in low phosphorus soil in the absence of the fungus.

(b) Mycorrhizal associations. Experimental seedlings were grown under the same conditions as the controls, except that the fungus was present. The seedlings formed mycorrhizal associations with the fungus.

RESULTS AND CONCLUSIONS: Growth of the plants that formed mycorrhizal associations was significantly enhanced. Mycorrhizal associations enhance the growth of western red cedar plants. Many similar studies using other types of plants and other types of soil have confirmed the importance of mycorrhizal association to plant growth.

FIGURE 29-19 The effect of mycorrhizae on western red cedar (Thuja plicata) seedlings

More recently, researchers have suggested that the lichen partnership is not really a case of mutualism but one of controlled parasitism of the photoautotroph by the fungus. Lichens typically exhibit one of three different growth forms (FIG. 29-20b). Crustose lichens are flat and grow tightly against their substrate (the surface they are growing on). Foliose lichens are also flat, but they have leaflike lobes and are not so tightly pressed to the substrate. Fruticose lichens grow erect and have many branches. Able to tolerate extremes of temperature and moisture, lichens grow in almost all terrestrial environments except polluted cities. They exist farther north than any plants of the arctic region and are equally at home in the steaming equatorial rain forest. They grow on tree bark, leaves, and exposed rock surfaces, from solidified lava to tombstones. In fact, lichens are often the first organisms to inhabit rocky areas, and their growth in these areas is important in forming soil from rock. They secrete acid that gradually etches tiny cracks in the rock, releasing minerals. This process sets the stage for further disintegration of the rock by wind and rain. When the lichens themselves die and are decomposed, they become part of the soil. Reindeer mosses of the arctic region, which serve as the main source of food for migrating herds of caribou, are actually lichens, not mosses. Some lichens produce colored pigments. One of them, orchil, is used to dye wool, and another, litmus, is widely used in chemistry laboratories as an acid–base (pH) indicator.

Lichens vary greatly in size. Some are almost invisible, whereas others, such as the reindeer mosses, may cover many square kilometers of land with an ankle-deep growth. Growth proceeds slowly; the radius of a lichen may increase by less than 1 mm each year. Some mature lichens are thought to be thousands of years old. Lichens absorb minerals from the air, rainwater, and the surface on which they grow. They cannot excrete the elements they absorb, and perhaps for this reason they are extremely sensitive to toxic compounds. This sensitivity was first reported in 1866 by a Finnish biologist who observed that lichens growing on tree trunks in Paris were poorly developed or sterile. He deduced that lichens could be used to measure air purity. Today, reduction in lichen growth is used as a sensitive indicator of air pollution, particularly from sulfur dioxide. In one study, investigators demonstrated a relationship between lung cancer and air pollution by comparing the locations of low lichen biodiversity (and therefore of air pollution) with the locations of lung cancer deaths in young males. The return of lichens to an area indicates an improvement in air quality. Lichens reproduce mainly by asexual means, usually by fragmentation, a process in which special dispersal units of the lichen, called soredia, break off and, if they land on a suitable surface, establish themselves as new lichens. Soredia contain cells of both partners. In some lichens, the fungus produces ascospores, which may be dispersed by wind and find an appropriate algal partner only by chance.

Fruticose lichen (Ramalina)

Soredia Surface layer (fungal hyphae) Fungal hyphae interwoven with photoautotroph Loosely woven hyphae Rock or other surface to which lichen is attached

Crustose lichens (Bacidia, Lecanora) Foliose lichen (Parmelia) Fred M. Rhoades

(a) This cross section of a typical lichen shows distinct layers. The soredium (pl., soredia), an asexual reproductive structure, consists of clusters of algal or cyanobacterial cells enclosed by fungal hyphae.

Bottom layer (fungal hyphae)

FIGURE 29-20 Animated Lichens These organisms are a combination of a fungus and an alga or cyanobacterium.

Review ■ ■ ■

What is the ecological importance of fungal decomposers? What is the importance of mycorrhizae? What is a lichen?

29.5 ECONOMIC, BIOLOGICAL, AND MEDICAL IMPACT OF FUNGI ■ ■ LEARNING OBJECTIVES 10 Summarize some specific ways that fungi impact humans economically. 11 Summarize the importance of fungi to biology and medicine; describe how fungi infect plants and humans, describing at least one fungal animal disease and one fungal plant disease.

The same powerful digestive enzymes that fungi use to decompose wastes and dead organisms can also be used with great efficiency to reduce wood, fiber, and food to their basic components. Many species of basidiomycetes have enzymes that break down the lignin in wood. (Lignin is the second most abundant organic compound on Earth, second only to cellulose.) From the human perspective,

(b) Lichens vary in color, shape, and overall appearance. Three growth forms—crustose, foliose, and fruticose—are shown on a maple branch in Washington State.

various fungi cause incalculable damage to stored goods and building materials each year. Bracket fungi, for example, cause enormous losses by decaying wood, both in living trees and in stored lumber. Some fungi cause serious diseases in animals and plants. Yet fungi also contribute to our quality of life. They are responsible for economic gains as well as losses. People eat them and grow them to make various medications, such as penicillin. We use them to make certain industrial chemicals and for bioremediation. Renewable fuel companies are actively searching for fungi and other microbes that can produce fuel. Recently, a fungus was discovered that synthesizes 55 hydrocarbons, perhaps to inhibit the growth of other organisms. Researchers may be able to use the genes of this fungus to engineer other microorganisms to more efficiently produce fuel.

Fungi provide beverages and food Humans exploit the ability of yeasts to produce bread and alcoholic beverages. Yeasts produce ethyl alcohol and carbon dioxide from glucose and other sugars by fermentation (see Chapter 8). Yeast species of the genus Saccharomyces (ascomycetes) are used to produce wine, beer, and other fermented beverages. Wine is produced when yeasts ferment fruit sugar, and beer results when yeasts ferment sugar derived from starch in grains (usually barley).

names as “destroying angel” (A. virosa) and “death cap” (A. phalloides). Eating a single mushroom of either species can be fatal. Certain species of mushrooms cause intoxication and hallucinations. The sacred mushrooms of the Aztecs—Conocybe and Psilocybe—are still used in religious ceremonies by native peoples of Central America for their hallucinogenic properties. The chemical ingredient psilocybin is responsible for the trances and visions experienced by those who eat these mushrooms. Ingestion of psychoactive mushrooms is dangerous because negative reactions vary considerably, from mild indigestion, sweating, and heart palpitations, to death. In addition, the possession and use of such mushrooms are illegal in the United States and some other countries.

Saccharomyces cerevisiae, referred to as baker’s yeast, is used to bake bread, pizza, and other wheat products. During the process of making bread, carbon dioxide produced by yeast becomes trapped in dough as bubbles, causing the dough to rise; this gives leavened bread its light quality. Both the carbon dioxide and the alcohol produced by the yeast escape during baking. The unique flavor of cheeses such as Roquefort, Brie, Gorgonzola, and Camembert is produced by species of Penicillium. For example, P. roquefortii, found in caves near the French village of Roquefort, is used to make Roquefort cheese. By French law, only cheeses produced in this area can be called Roquefort cheese. (The blue spots in Roquefort and certain other cheeses are masses of conidia.) Aspergillus tamarii and certain other fungi are used to produce soy sauce by fermenting soybeans with the fungi for at least three months. (In the United States, soy sauce is often made by adding flavoring to salt water rather than by soaking fermented soybeans.) Soy sauce enriches other foods with more than just its special flavor. It also adds vital amino acids from both the soybeans and the fungi themselves, which, in some parts of the world, supplement a low-protein rice diet. Among the basidiomycetes, there are some 200 kinds of edible mushrooms and about 70 species of poisonous ones. Many edible mushrooms are cultivated commercially. The mushroom Agaricus brunnescens is the principal fungal species grown extensively for food. About 30 other mushroom species, such as oyster, shiitake, portobello, and straw mushrooms, are available in supermarkets. Morels, which superficially resemble mushrooms, and truffles, which produce underground fruiting bodies, are ascomycetes (FIG. 29-21). Truffles are now cultivated as mycorrhizal fungi on the roots of tree seedlings. Edible and poisonous mushrooms can look very much alike and may even belong to the same genus. There is no simple way to tell them apart; an expert must identify them. Some of the most poisonous mushrooms belong to the genus Amanita (FIG. 29-22). Toxic species of this genus have been appropriately called such

Fungi are important to modern biology and medicine

(a) The yellow morel (Morchella esculenta). This morel grows 6 to 10 cm (2.5 to 4 in) tall. It is found throughout North America. Photographed in Michigan.

FIGURE 29-21 Edible ascomycetes

John D. Cunningham /Visuals Unlimited

Richard Shiell/Dembinsky Photo Associates

The yeast Saccharomyces cerevisiae has served as a model eukaryotic cell (see Fig. 17-6a). It was the first eukaryote whose genome was sequenced, and with its 6000 genes, it has the smallest genome of any eukaryotic model organism. Molecular biologists are in the process of determining the functions of the proteins encoded by its genes. Biologists have used S. cerevisiae to study molecular genetics, including how genes regulate cell division. Researchers continue to use this yeast to study such problems as genetic recombination, and the correlation between cell age and cancer. Saccharomyces cerevisiae is also being used to study the mechanism of action of antifungal drugs and resistance to these drugs. Biologists have used the ascomycete Aspergillus nidulans, an opportunistic pathogen of humans, to study mitosis and other cell processes. This fungus has provided valuable knowledge about the genetics of microtubules. Biologists are using recombinant DNA techniques to manipulate yeasts and certain filamentous fungi to produce important biological molecules, such as hormones. Among the many genes that have been cloned in yeast are those for insulin, human growth hormone, and molecules important in

(b) The Oregon white truffle (Tuber gibbosum). The truffle, which is found underground near Douglas firs and possibly oak trees in British Columbia and northern California, is 1 to 5 cm (0.4 to 2 in) wide. People find these subterranean ascocarps with the help of trained dogs or pigs. Here, truffles are shown whole and sectioned to show the conspicuous white, marbled tissue.

James W. Richardson/CBR Images

Fungi are used in bioremediation and to biologically control pests

FIGURE 29-22 Poisonous mushrooms The destroying angel (Amanita virosa) is an extremely poisonous mushroom that is distinguished, as are other amanitas, by the ring of tissue around its stalk and by the underground cup from which the stalk protrudes. About 50 g (2 oz) of this mushroom can kill an adult man. The destroying angel, which is 7.5 to 20 cm (3 to 8 in) tall, is found in grass or near trees throughout North America.

immune function. These procedures allow researchers to produce unlimited amounts of these compounds for study and eventual medical use. Fungi produce useful drugs and chemicals. Discovered in 1928 by the British bacteriologist Alexander Fleming, penicillin, produced by the mold Penicillium notatum, is still among the most widely used and effective antibiotics (see Chapter 1). Other drugs derived from fungi include the cephalosporin antibiotics (produced by Cephalosporium), statins (used to lower blood cholesterol levels), and cyclosporine (used to suppress immune responses in patients who receive organ transplants). Fumagillin, a chemical produced by the ascomycete Aspergillus fumigatus, inhibits the formation of new blood vessels. Because solid tumors need a rich blood supply, fumagillin shows promise as an anticancer agent. Fumagillin is also used to treat diseases caused by microsporidia. Researchers have identified several other promising compounds produced by fungi that are antiviral or that destroy cancer cells. The ascomycete Claviceps purpurea infects the flowers of rye plants and other cereals. It produces a structure called an ergot where a seed would normally form in the grain head. When livestock eat this grain or when humans eat bread made from ergotcontaminated rye flour, they may be poisoned by the extremely toxic substances in the ergot. However, some ergot compounds are now used clinically in small quantities as drugs to induce labor, to stop uterine bleeding, to treat high blood pressure, and to relieve one type of migraine headache.

Some fungi can biodegrade pesticides, herbicides, coal tars, and petroleum. Fungi convert these products into carbon dioxide and the basic elements of which they are composed. These fungi can be used along with certain bacteria to decontaminate farm land and to clean up oil spills. Researchers are investigating fungi—for example, certain species of microsporidia—for the biological control of pathogens and insect pests. Some of these species are already being used to parasitize insect pests. In some cases, they interfere with reproduction in their insect host. Other microsporidia completely control the metabolism and reproduction of the host. Researchers are studying the use of microsporidia in controlling the spread of malaria. A recent study showed that when female Anopheles mosquitoes are infected with microsporidia, their blood feeding is decreased. Fungal infection of the mosquito also interferes with development of Plasmodium, the protist that causes malaria.

Some fungi cause diseases in humans and other animals Certain ascomycetes cause superficial infections in which only the skin, hair, or nails are infected. Ringworm, athlete’s foot, and jock itch are examples of superficial fungal infections. Because these fungi infect dead layers of skin that are not fed by capillaries, the immune system cannot launch an effective response. Many pathogenic fungi are opportunists that cause infections only when the body’s immune system is compromised, for example, in patients with HIV. Cancer patients and organ transplant recipients who are given medication to suppress their immune systems are also at risk. Candida is an ascomycete that inhabits the human mouth and vagina. The immune system and the normal bacteria of these regions normally prevent this yeast from causing infection. However, when the immune system is compromised, Candida multiplies, causing thrush, a painful yeast infection of the mouth, throat, and vagina. The ascomycete Aspergillus fumigatus is usually harmless but causes aspergillosis in people with lowered immune function. During the course of aspergillosis, the fungus can invade the lungs, heart, brain, kidneys, and other vital organs and cause death. Other fungi infect internal tissues and organs and may spread through many regions of the body. Histoplasmosis, for example, is a lung infection caused by inhaling spores of a fungus common in soil contaminated with bird feces. Most people in the eastern and midwestern parts of the United States have been exposed to this fungus at some time, and an estimated 40 million Americans have had mild infections. Fortunately, the infection is usually confined to the lungs and is of short duration, but if the infection spreads through the blood to the heart, brain, or other parts of the body, it can be serious and sometimes fatal. Some fungi produce poisonous compounds collectively called mycotoxins. A few species of Aspergillus, for example, produce potent mycotoxins called aflatoxins that harm the liver and are known carcinogens. Foods on which aflatoxin-producing fungi commonly grow include peanuts, pecans, corn, and other grains. Other foods

that may contain traces of aflatoxins include animal products such as milk, eggs, and meat (from animals that consumed feed contaminated by aflatoxin). Avoiding aflatoxin in the diet is impossible, but exposure should be minimized as much as possible. Any human food or animal forage product that has become moldy should be suspected of aflatoxin contamination and should be discarded. Fungi contribute to sick-building syndrome, a situation in which occupants of a building experience acute adverse health effects linked to the time they spend in a given building. Moldrelated insurance claims amount to hundreds of millions of dollars each year. When conditions are moist, molds can grow on carpets, leather, cloth, wood, insulation, and food. Mold spores, fragments, and mold products make their way into the air, and people are exposed through inhalation as well as by skin contact. Exposure to molds and their toxins has been linked to depressed immune function, irritation of the throat and respiratory passageways, infection, and toxicity. The most common responses to mold exposure are allergic reactions that range from mild to severe illnesses, including hay fever, sinusitis, asthma, and dermatitis.

Fungi cause many important plant diseases

(a) Brown rot of peaches. This disease is caused by Monilinia fruticola, an ascomycete. Photographed in Oregon.

FIGURE 29-24 Fungi that cause plant diseases

Hypha

Epidermis Stoma Airspace

Leaf

Haustoria

FIGURE 29-23 How a fungus parasitizes a plant In this example, the hypha enters the leaf through a stoma. The hypha grows, branching extensively through the internal air spaces, and penetrates plant cells with specialized hyphal extensions called haustoria.

A plant often becomes infected after hyphae enter through stomata (pores) in the leaf or stem or through wounds in the plant body (FIG. 29-23). Alternatively, the fungus may produce cutinase, an enzyme that dissolves the waxy cuticle that covers the surface of leaves and stems. After dissolving the cuticle, the fungus easily invades the plant tissues. As the mycelium grows, it may stay mainly between the plant cells or it may penetrate the cells. Parasitic fungi often produce special hyphal branches called haustoria (sing., haustorium) that penetrate the host cells and obtain nourishment from the cytoplasm. Ascomycetes cause serious plant diseases, including powdery mildew, chestnut blight, Dutch elm disease, apple scab, wilt on potatoes, and brown rot, which attacks cherries, peaches, plums, and

© Nigel Cattlin /Photo Researchers, Inc.

Runk /Schoenberger, from Grant Heilman

Kathy Merrifield / Photo Researchers, Inc.

Fungi are more destructive to plants than any other diseasecausing organism. They are responsible for about 70% of all major crop diseases. Fungi cause serious epidemic diseases that spread rapidly and often result in complete crop failure. Fungal plant diseases costs billions of dollars in agricultural damage yearly. All plants are apparently susceptible to some fungal infection. Damage may be localized in certain tissues or structures of the plant, or the disease may be systemic and spread throughout the entire plant. Fungal infections may cause stunting of plant parts or of the plant; they may cause wartlike growths or kill the plant.

Spore

(b) Corn smut on an ear of sweet corn. This fungal disease is caused by Ustilago maydis, a basidiomycete.

(c) Black stem rust infection on wheat (Triticum sp.) stem. This plant disease is caused by Puccinia graminis, a basidiomycete.

apricots (FIG. 29-24a). Basidiomycetes cause smuts and rusts that attack corn, wheat, oats, and other grains (FIG. 29-24b and c). Some fungal parasites, such as the basidiomycete (Puccinia graminis) that causes black stem rust of wheat, have complex life cycles that involve two or more different host plants and the production of several kinds of spores. Before the late 1950s, black stem rust outbreaks occurred every few years somewhere in the world, destroying entire wheat crops. By the early 1960s, Nobel prize winner Norman Borlaug and other researchers had developed rust-resistant varieties of wheat, heralding in the green revolution. However, new mutations have put Puccinia and wheat rust back in the news. The new strain has spread through parts of Africa and into the Middle East. Each fungus releases billions of spores, which the wind can blow for hundreds of miles, spreading the fun-

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gus to new regions of the world. Its spread may lead to widespread food shortages. The Puccinia genome has recently been sequenced, and researchers are developing methods to quickly diagnose the disease. They are also working to develop new varieties of diseaseresistant wheat.

Review ■

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Some dictionaries erroneously define a morel as a type of mushroom. Why is this definition incorrect? How are fungi important in modern biology and medicine? What are three important fungal diseases of humans? Why is the mutated strain of the fungus that causes wheat rust of serious concern?

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

In most fungi, perforated septa, or cross walls, divide the hyphae into individual cells. In some fungi, the hyphae are coenocytic, that is, they form an elongated, multinuclear cell.

Chytrids

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29.2 (page 603) 3 Describe the life cycle of a typical fungus, including sexual and asexual reproduction. ■ Most fungi reproduce both sexually and asexually by means of spores. Spores are produced on aerial hyphae. When fungal spores land in a suitable spot, they germinate. ■ When fungi of two different mating types meet, their hyphae fuse, a process called plasmogamy. The cytoplasm fuses, but the nuclei remain separate. The fungi enter a dikaryotic (n + n) stage in which each new cell formed has one nucleus of each type. ■ Karyogamy, fusion of the nuclei, takes place in the hyphal tip and results in a diploid (2n) zygote nucleus. ■ Meiosis produces four genetically different haploid (n) nuclei. Each nucleus becomes part of a spore. When the spores germinate, they form new mycelia by mitosis. ■ Genetically similar asexual spores are produced by mitosis. When these spores germinate, they also develop into mycelia.

29.3 (page 605) 4 Give arguments to support the hypothesis that fungi are opisthokonts, more closely related to animals than to plants. ■ Like animals, some fungi have flagellate cells—for example, chytrid gametes and spores; the flagellate cells propel themselves with a single posterior flagellum. Also like animal cells, fungal mitochondria have platelike cristae.

Basidiomycetes

Learn more about mycelia by clicking on the figure in CengageNOW.

Ascomycetes

Fungi are eukaryotic heterotrophs that secrete digestive enzymes onto their food source and then absorb the predigested food. Fungi are characterized by cell walls that contain chitin. 2 Describe the body plan of a fungus. ■ A fungus may be a unicellular yeast or a filamentous, multicellular mold. The body of most multicellular fungi consists of long, threadlike filaments called hyphae that branch and form a tangled mass called a mycelium. ■

Glomeromycetes

1 Describe the distinguishing characteristics of fungi.

Based on chemical and structural characters, fungi are classified, along with animals and choanoflagellates, as opisthokonts. 5 Give arguments to support the hypothesis that chytrids may have been the earliest fungal group to evolve from the most recent common ancestor of fungi. ■ Chytrids, or chytridiomycetes, produce flagellate spores at some stage in their life cycle. No other fungi have flagella. Thus, chytrids probably were the earliest fungi to evolve; the most recent common ancestor of all fungi was a flagellate protist. 6 List distinguishing characteristics, describe a typical life cycle, and give examples of each of the following fungal groups: chytridiomycetes, zygomycetes, glomeromycetes, ascomycetes, and basidiomycetes. ■

Zygomycetes

29.1 (page 602)

Evolution of ascospores Evolution of basidiospores Evolution of dikaryotic stage

Loss of flagellum Flagellate common ancestor ■

Chytrids reproduce both asexually and sexually. Their gametes and zoospores are flagellate. Allomyces, a common chytrid, spends part of its life as a multicellular haploid thallus and part as a multicellular diploid thallus. The haploid thallus produces two types of flagellate gametes that fuse. Both plasmogamy and karyogamy occur, producing a flagellate zygote. The diploid thallus bears zoosporangia that produce diploid zoospores and resting sporangia in which haploid zoospores form by meiosis. The haploid zoospores form new haploid thalli.

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Zygomycetes, such as the black bread mold, Rhizopus, form a haploid thallus that produces both asexual spores and sexual spores. Asexual spores germinate and form new thalli. In sexual reproduction, hyphae of two different haploid mating types form gametangia. Plasmogamy occurs as the gametangia fuse. Karyogamy occurs, and a diploid zygote is formed; the zygote develops into a zygospore. Meiosis produces recombinant haploid zygospores. When zygospores germinate, each hypha develops a sporangium at its tip. Spores are released and develop into new hyphae. Microsporidia, currently classified as zygomycetes, are opportunistic pathogens that penetrate and infect animal cells with their long, threadlike polar tubes. Glomeromycetes have coenocytic hyphae. They reproduce asexually with large, multinucleate spores called blastospores. Glomeromycetes are symbionts that form intracellular associations called mycorrhizae with the roots of plants. Because they extend their hyphae into root cells, glomeromycetes are endomycorrhizal fungi. The most common endomycorrhizae are called arbuscular mycorrhizae because the hyphae inside the root cells form branched, tree-shaped structures known as arbuscules. Ascomycetes include yeasts, cup fungi, morels, truffles, and bluegreen, pink and brown molds. Some ascomycetes form mycorrhizae; others form lichens. Ascomycetes produce asexual spores called conidia; they produce sexual spores called ascospores in saclike asci. The asci line a fruiting body called an ascocarp. In ascomycetes, haploid mycelia of opposite mating types produce septate hyphae. Plasmogamy occurs, and nuclei are exchanged. A dikaryotic n + n stage occurs in which hyphae form and produce asci and an ascocarp. Karyogamy occurs, followed by meiosis. The recombinant nuclei divide by mitosis, producing eight haploid nuclei that develop into ascospores. When the ascospores germinate, they can form new mycelia. Basidiomycetes include mushrooms, puffballs, bracket fungi, rusts, and smuts. These fungi produce sexual spores called basidiospores on the outside of a basidium. Basidia develop on the surface of gills in mushrooms; mushrooms are a type of basidiocarp (a fruiting body). Hyphae in the basidiomycetes have septa. Plasmogamy occurs with the fusion of two hyphae of different mating types. A dikaryotic secondary mycelium forms. Then a basidiocarp develops, and basidia form. Karyogamy occurs, producing a diploid zygote nucleus. Meiosis produces four haploid nuclei that become basidiospores. When basidiospores germinate, they form haploid primary mycelia. Explore fungus life cycles by clicking on the figures in CengageNOW.

29.4 (page 615) 7 Summarize the ecological significance of fungi as decomposers. Most fungi are decomposers that break down organic compounds in dead organisms, leaves, garbage, and wastes into simpler nutrients that can be recycled. 8 Describe the important ecological role of mycorrhizae. ■ Mycorrhizae are mutualistic associations between fungi and the roots of plants. The fungus supplies water and nutrient minerals to the plant; the fungus obtains organic compounds from the plant. Glomeromycetes form endomycorrhizae with roots. Ascomycetes and basidiomycetes form ectomycorrhizae with tree roots; they do not penetrate the root cells. 9 Characterize the unique nature of a lichen. ■ A lichen is a combination of a fungus and a photoautotroph (an alga or cyanobacterium). In this symbiotic relationship, the photoautotroph provides the fungus with organic compounds. The fungus may provide the photoautotroph with shelter, water, and minerals. Lichens have three main forms: crustose, foliose, and fruticose. ■

Learn more about lichens by clicking on the figure in CengageNOW.

29.5 (page 619) 10 Summarize some specific ways that fungi impact humans economically. Fungi cause huge economic losses by damaging food and crops. On the other hand, some fungi, like mushrooms, are foods; others, like yeasts, are used to produce beer, wine, and bread, and still others are used to produce cheeses or industrial chemicals. 11 Summarize the importance of fungi to biology and medicine; describe how fungi infect plants and humans, describing at least one fungal animal disease and one fungal plant disease. ■ Biologists use the yeast Saccharomyces cerevisiae and other fungi as model organisms for research in molecular biology and genetics. Fungi are also being investigated for the biological control of insects, such as the mosquitoes that transmit malaria. ■ Fungi are used to make many medications, including penicillin and other antibiotics; they are used in bioremediation and to biologically control pests. ■ Fungi are opportunistic pathogens in humans. They cause human diseases, such as histoplasmosis; some fungi produce mycotoxins, such as aflatoxins, which can cause liver damage and cancer. ■ Fungal hyphae infect plants through stomata. Hyphal branches called haustoria penetrate plant cells and obtain nourishment from the cytoplasm. Fungi cause many important plant diseases, including brown rot, corn smut, and wheat rust. ■

T E S T YO U R U N D E R S TA N D I N G 1. Fungi are (a) eukaryotes and opisthokonts (b) prokaryotes and opisthokonts (c) flagellate and dikaryotic (d) autotrophic eukaryotes (e) heterotrophs with cellulose cell walls 2. Which of the following fungi does not have a mycelium? (a) black bread mold (b) yeast (c) decomposer cup fungus (d) cultivated mushroom (e) Penicillium 3. A cell described as n + n is (a) monokaryotic (b) diploid (c) haploid (d) coenocytic (e) dikaryotic 4. With the exception of chytridiomycetes, fungi are generally disseminated by (a) water currents (b) fragmentation of hyphae (c) soredia (d) airborne spores (e) flagellate zoospores

5. Which statement is not true of the chytrids? (a) they are simple aquatic fungi (b) they produce motile cells with single, posterior flagella (c) they undergo both sexual and asexual reproduction (d) their cells are dikaryotic (e) they were the earliest fungi to evolve 6. Which statement is not true of the zygomycetes? (a) many members of this group form endomycorrhizae with tree roots (b) their sexual spores are called zygospores (c) they undergo both sexual and asexual reproduction (d) plasmogamy and karyogamy occur (e) they have coenocytic hyphae

7. Glomeromycetes (a) reproduce mainly by sexual spores called glomerospores (b) are characterized by unique structures called polar tubes (c) associate with cyanobacteria to form lichens (d) include many opportunistic pathogens that cause human disease (e) form arbuscular endomycorrhizae with tree roots 8. The ascomycete life cycle typically includes (a) mainly diploid thalli (b) the formation of a thick zygosporangium (c) the production of eight haploid ascospores within an ascus (d) the production of microsporidia (e) the production of ascospores, zoospores, and conidia at different stages 9. Which statement is not true of the basidiomycetes? (a) they produce a secondary mycelium with n + n hyphae (b) their sexual spores are called basidiospores (c) they have a diploid thallus that

produces zoospores (d) reproduction is mainly sexual (e) they have microscopic basidia 10. A combination organism consisting of a photoautotroph and a fungus is called _____ (a) an arbuscular endomycorrhiza (b) an ectomycorrhiza (c) a lichen (d) a pathogenic agent (e) an aflatoxin 11. Mutualistic associations between fungi and the roots of plants are called (a) lichens (b) mycorrhizae (c) pathogenic associations (d) parasitic haustoria (e) mycotoxic symbioses 12. When a fungus infects a plant, it (a) infiltrates leaves with lichens (b) forms relationships by attaching mycorrhizae to stems (c) secretes powerful digestive juices onto the leaves (d) uses haustoria to dissolve roots (e) enters leaves or stems through stomata

CRITICAL THINKING 1. Explain the statement “Mushrooms are like the tips of icebergs.” If you do not see mushrooms in your lawn, can you conclude that no fungi live there? Why or why not?

4. The secondary mycelium of a basidiomycete is shown here. (a) How would you describe its cells? (b) What stage of its life cycle is it in? (c) Draw the next steps in this stage of its life cycle.

2. EVOLUTION LINK. Justify (a) classifying fungi as opisthokonts; (b) classifying microsporidia as fungi; (c) grouping ascomycetes and basidiomycetes as sister clades. 3. SCIENCE, TECHNOLOGY, AND SOCIETY. Biologists have discovered that many mycorrhizal fungi are sensitive to a low pH. What human-caused environmental problem may prove catastrophic for these fungi? How may this problem affect their plant partners? What measure could we take to decrease the problem? Additional questions are available in CengageNOW at www.cengage.com/ login.

30

An Introduction to Animal Diversity

Charles V. Angelo/Photo Researchers, Inc.

The tube sponge (Callyspongia vaginalis). This animal, sometimes mistaken for a plant, ranges in color from purple to blue to gray. It is common on coral reefs in the Caribbean, from Florida to Mexico.

KEY CONCEPTS

A

30.1 Although they are a diverse group, animals share

thought sponges were plants because they did not move from place to

many characters. They are multicellular, eukaryotic heterotrophs composed of cells specialized to perform specific functions. Most animals are diploid organisms that reproduce sexually, and most have a nervous and muscular system.

place. Some people still mistake certain marine animals, such as sponges

30.2 Animals evolved in marine environments, and members of most animal phyla still inhabit marine environments. However, many animals are adapted to life in fresh water and others to terrestrial habitats.

Biologists have described and named more than 1.5 million species of

30.3 The common ancestors of animals are choanoflagellates; choanoflagellates, fungi, and animals are a monophyletic group known as opisthokonts.

lthough members of most animal species are readily recognizable as animals, the identity of some others is less obvious. Early naturalists

and corals, for plants (see photograph). Locomotion is not a requirement for being classified as an animal. Animal phylogeny is an exciting and rapidly changing field of study. animals, and 15,000 to 20,000 new species are named each year. Millions more probably remain to be discovered and classified. Interestingly, an estimated 99% of all animal species that ever inhabited our planet are extinct. Taxonomists have assigned the extant (living) animals to about 35 phyla. Molecular studies have confirmed that many of these groups are

30.4 Biologists classify animals based on many character-

monophyletic, that is they consist of all of the descendants and only the

istics, including their morphology, features of their early development, and on molecular data; they generally agree that bilateral animals split into at least three major clades.

descendants of a common ancestor. (Recall from Chapter 23 that a monophyletic group is called a clade.)

Animal groups that are not monophyletic have been split or reorganized, with some members of the groups being reassigned to other taxa. It is important to remember that the classification of animals and the relative positions of animal groups are a work in progress. As they consider new data, systematists redraw the tree of animal life. In this chapter we discuss the characters of animals and their habitats. We then explore animal origins and some of the criteria biologists use to determine evolutionary relationships and to classify animals. In Chapter 31 we describe three phyla traditionally viewed as diverging early in the evolutionary history of animals. Then we discuss one of the major clades of animals: the protostomes. (In Chapter 32 we focus on the deuterostomes, which include the echinoderms and chordates—the clade to which we belong.) Many hypotheses are presented in these chapters, and we will discuss

6. Most animals have nervous systems and muscle systems that enable them to respond rapidly to stimuli in their environment. 7. Most animals are diploid organisms that reproduce sexually, with large, nonmotile eggs and small, flagellate sperm. A haploid sperm unites with a haploid egg, forming a diploid zygote (fertilized egg). 8. Animals go through a period of embryonic development. The zygote undergoes cleavage, a series of mitotic cell divisions. During cleavage the zygote develops into a hollow ball of cells called a blastula. Although some animals develop directly into adults, the majority first develop into a larva, a sexually immature form that may look very different from the adult (FIG. 30-1). The larva differs from the adult in many ways, including where it lives (its habitat), how it moves, and what it eats. Larvae typically go through metamorphosis, a developmental process that converts the immature animal into a juvenile form that can then grow into an adult.

many examples of how systematists revise the relationships of the branches of the animal phylogenetic tree in response to new data.

Review ■

30.1 ANIMAL CHARACTERS ■ ■ LEARNING OBJECTIVE 1

Describe several characteristics common to most animals.

Animals are so diverse that for almost any definition we can find exceptions. We can best describe animals by the characteristics they share: 1. Animals are multicellular eukaryotes. In contrast to plants, algae, and fungi, their cells lack cell walls. Instead, structural support depends on collagen and other structural proteins. Collagen is an important shared derived character in animals (see Chapter 23). 2. Animals are heterotrophs. As consumers, they depend on producers for their raw materials and energy. In contrast to the fungi, most animals ingest their food first and then digest it inside the body, usually within a digestive system. 3. Cells that make up the animal body are specialized to perform specific functions. In all but the simplest animals, cells are organized to form tissues, and tissues are organized to form organs. In small animals with simple body plans, life processes such as gas exchange, circulation of materials, and waste disposal can take place by diffusion of gases and other substances directly to and from the environment. Specialized organ systems perform these functions in large animals. 4. Animals have diverse body plans. The term body plan refers to the basic structure and functional design of the body. An animal’s body plan and lifestyle are adapted to its methods of obtaining food and reproducing. 5. Most animals are capable of locomotion at some time during their life cycle. Some animals (such as sponges and corals) move about as larvae (immature forms) but are sessile (firmly attached to the ground or some other surface) as adults (see chapter opening photograph).

For centuries, scientists classified sponges as plants, but now they are classified as animals. What criteria do biologists use to classify them as animals?

30.2 ADAPTATIONS TO OCEAN, FRESHWATER, AND TERRESTRIAL HABITATS ■ ■ LEARNING OBJECTIVE 2

Compare the advantages and disadvantages of life in the ocean, in fresh water, and on land.

Fossil evidence suggests that animals evolved in shallow, marine environments during the Proterozoic eon, at least 600 million years ago (mya; see Chapter 21). Although animals are now distributed in virtually every environment, at least some members of most animal phyla still inhabit marine environments.

Marine habitats offer many advantages The buoyancy of sea water provides support, and its large volume keeps the water temperature relatively stable. The body fluids of most invertebrates have about the same osmotic concentration as sea water, so fluid and salt balance are more easily maintained than in fresh water. Plankton, which consists of the mainly microscopic animals and protists that are suspended in water and float with its movement, provides a ready source of food for many aquatic animals. Life in the ocean also presents some challenges. Although the continuous motion of water brings nutrients to animals and washes their wastes away, animals must be able to cope with the water’s movements and the currents that could sweep them away. Squids, fishes, and marine mammals have evolved as strong swimmers, usually able to direct their movements and maintain their location. However, most invertebrates and young

Dr. Magnus Kjaergaard

Doug Perrine / SeaPics.com

(a) Spiny lobster larva. This larva is hitching a ride on a jellyfish. Over a period of months, the larva passes through several stages before becoming an adult. Photographed in open ocean at night, Hawaii.

(b) Adult spiny lobster. The adult has two long antennae used to sense movement. Spiny lobsters lack large claws. The spines on their back help protect them. In this image you can see the two large spines above the animal’s eyes. Spiny lobsters leave their hiding places at night to hunt.

FIGURE 30-1 Larva and adult spiny lobster (Panulirus sp.) Most animals go through a larva stage before developing into an adult. The larva usually differs from the adult in size, appearance, and in lifestyle.

vertebrates cannot swim strongly, and they have adapted in various ways to the tides and currents. Some sessile animals attach permanently to a stable structure such as a rock. Others burrow in the sand and silt that cover the sea bottom. Many invertebrates have adapted by maintaining a small body size and becoming part of the plankton. As they are tossed about, their food supply continues to surround them.

Some animals are adapted to freshwater habitats Far fewer kinds of animals make their homes in fresh water than in the ocean because living in this habitat is more difficult. Fresh water is hypotonic to the tissue fluids of animals, so water tends to move into the animal by osmosis. To survive in this habitat, freshwater species must have mechanisms for removing excess water while retaining salts. This osmoregulation requires an expenditure of energy. Fresh water offers a much less constant environment than sea water. Animals that inhabit fresh water must have adaptations for surviving variations in oxygen content, temperature, turbidity (because of sediments suspended in the water), and even water volume. In addition, fresh water generally contains less food than the sea.

Terrestrial living requires major adaptations Living on land is even more difficult than living in fresh water, and the evolution of terrestrial animals involved major adaptations. Analyzing the fossil record, many biologists hypothesize that the

first air-breathing terrestrial animals were scorpion-like arthropods that came ashore in the Silurian period about 444 mya. The first vertebrates to inhabit terrestrial environments, the amphibians, did not appear until the Devonian period, about 30 million years later. The chief problem facing all terrestrial organisms is desiccation (drying out). Water is constantly lost by evaporation and is often difficult to replace. A body covering adapted to minimize fluid loss helps solve this problem in many terrestrial animals (FIG. 30-2). Location of the respiratory surface deep within the animal also helps prevent fluid loss. Thus, the gills of aquatic animals are typically located externally, but lungs and tracheal tubes of terrestrial animals are typically found deep within the body. Reproduction on land also poses challenges to protect gametes and the developing offspring from desiccation. Aquatic animals typically shed their gametes in the water, where fertilization occurs. The surrounding water also serves as an effective shock absorber that protects the delicate embryos as they develop. Some land animals, including most amphibians, return to the water for reproduction, and their larval forms develop in the water. The evolution of internal fertilization has permitted many terrestrial animals, including earthworms, land snails, insects, reptiles, birds, and mammals, to meet the desiccation challenge. Because these terrestrial animals transfer sperm from the body of the male directly into the body of the female by copulation, a watery medium continuously surrounds the sperm. Another important adaptation to reproduction on land is the tough, protective shell that surrounds the eggs of many species (see Fig. 30-2). Secreted by the female, this shell protects the developing embryo from drying out.

E. R. Degginger/Photo Researchers, Inc.

FIGURE 30-2 Adaptations to terrestrial life The tough, horny skin of the green iguana (Iguana iguana) has scales and is water resistant. Leathery eggs protect the embryos from drying out.

An alternative adaptation for terrestrial reproduction is development of the embryo within the moist body of the mother. Water has buoyancy that helps support animals that inhabit this environment. Air is less dense than water, and to inhabit the land, animals must have structures, such as a skeletal system and muscles, that support the body. The temperature extremes of terrestrial habitats also present challenges. In later chapters we discuss behavioral and physiological adaptations for maintaining body temperature.

Review ■

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What are some advantages of marine environments over freshwater and terrestrial habitats? What are some animal adaptations to the terrestrial environment?

30.3 ANIMAL EVOLUTION ■ ■ LEARNING OBJECTIVE 3

Use current hypotheses to trace the early evolution of animals.

Biologists generally agree that animals share a common ancestor with a group of protists known as choanoflagellates (see Fig. 26-21). The cells of these colonial flagellates became specialized to perform specific functions, such as movement, feeding, or reproduction. As this division of labor evolved, a colony of flagellates reached the level of cooperation and coordination that qualified it to be considered a single organism—the first animal. The choanoflagellates, fungi, and animals are a monophyletic group known as opisthokonts. Recall from Chapter 26 that opisthokonts are characterized by a posterior flagellum on motile cells. Historically, biologists depended on fossils, on similarities in body plan, and on patterns of development to determine evolutionary relationships among various groups of animals. Molecu-

lar systematics, the science that focuses on molecular structure to clarify evolutionary relationships, has provided additional data that are critical in answering questions about phylogeny. In many cases, molecular data have confirmed hypotheses that were based on morphology. Complex genomes were apparently present early during animal evolution. Molecular studies suggest that the ancestor of animals had more than 1500 genes not found in other eukaryotes. Some of these genes may be traced to horizontal gene transfer from other domains, followed by modification of the genes. Complex genomes have been described in the sea anemone and in other animals that are relatively simple morphologically. Molecular analyses indicate that the structure of genes that control development, RNA molecules, and many other molecules are very similar among all animal groups that have been studied. According to the principle of parsimony, such complex molecules are unlikely to have evolved multiple times (see Chapter 23). Thus, these data support the hypothesis that animals evolved only once. Animals are a monophyletic group.

Molecular systematics helps biologists interpret the fossil record The evolutionary history of animals has been vigorously debated, because early animals were soft-bodied forms that left few fossils. The scarcity of fossils has made it difficult to determine the age, rate of divergence, and number of branches of animal groups. In 2009, a research team found fossil traces in an oil field on the Arabian Peninsula that are thought to date back more than 635 million years. The fossil traces are steroids found only in the skeletal structures of certain sponges (demosponges). Before this discovery, the earliest known animal fossils were the Ediacaran biota from the Ediacaran period (600 mya to 542 mya). These fossils of small, simple animals suggest that sponges, jellyfish, and comb jellies were present during this period (see Fig. 21-9). Paleontologists have discovered many large, complex animal fossils in Chengjiang, an Early Cambrian (542 mya to 520 mya) fossil site in China, and in the Burgess Shale in British Columbia, a Middle Cambrian fossil site (520 mya to 515 mya). Fossils of most extant phyla (and also many extinct animals) have been found at these sites. The rapid appearance of an amazing variety of body plans during this time is known as the Cambrian radiation, or less formally as the Cambrian explosion (see Fig. 21-10). According to the Cambrian radiation hypothesis, which is based on the fossil record, major modifications in body plan that occurred during this time account for many branches of the animal tree. Studies of large molecular data sets suggest that most animal clades actually diverged over a very long period during the Proterozoic eon (2.5 bya to 542 mya). Thus, the animal phyla that first left fossils during the Cambrian radiation may have evolved several hundred million years before they appear in the fossil record. Biologists estimate that certain groups are about twice as old as the oldest fossils found to date. According to this view, the Cambrian radiation was a rapid evolution of new animal body plans among clades that already existed. Perhaps fossils of these early animals remain to be discovered in Proterozoic rocks. Another hypothesis

holds that a change in environmental conditions that occurred prior to the Cambrian radiation allowed fossils to form.

Biologists develop hypotheses about the evolution of development Changes in animal body plans are linked to changes in patterns of embryonic development. Biologists have long used similarities and differences in embryonic development to hypothesize how animal groups are related. Traditionally, biologists depended mainly on structural changes to compare the process of development in various groups. During the past 20 years, researchers have begun to study the molecular basis of developmental processes. They have identified the genes that direct the early development of the body plan and have discovered that many of these genes have been conserved during animal evolution. The same basic set of genes controls early development in all animal groups. Furthermore, the same genes are used in similar fashion to regulate development. Evolutionary developmental biology, sometimes referred to as Evo Devo, has become an important approach to studying animal relationships. Biologists compare molecular events, such as gene regulation during development, in various animal groups. Recall from Chapter 17 that Hox genes are a group of regulatory genes that specify the anterior-posterior axis during development (see Figure 17-13). The presence and number of Hox genes provide insights about evolutionary relationships. These genes have been identified in all of the bilateral animal groups that have been studied, suggesting that the last common ancestor of all bilateral animals had similar Hox genes. Recently, Hox genes were identified in a sea anemone (Nematostella vectensis), which is a cnidarian. Cnidarians are marine animals with radial symmetry, such as jellyfish. This finding suggests that the cnidarians share a common ancestor with the bilateral animals. Investigators think that all the Hox gene groups had evolved by the beginning of the Cambrian period. Mutations in Hox genes could have resulted in rapid changes in animal body plans. For example, regulation by Hox gene groups has been linked with the development of wings or legs. Similarities in molecular development among different animal groups suggest that they had a common ancestor.

Review ■ ■

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What was the Cambrian radiation? According to the current hypothesis, when did most major groups of animals evolve? How has the discovery of Hox genes helped biologists understand animal evolution?

30.4 RECONSTRUCTING ANIMAL PHYLOGENY ■ ■ LEARNING OBJECTIVES 4 Describe how biologists use morphology (including variations in body symmetry, number of tissue layers, and type of body cavity) and patterns of early development to infer relationships among animal phyla.

5

Cite specific examples of how data from molecular systematics have confirmed or modified traditional animal phylogeny. (Include identification of three major clades of bilateral animals.)

The basic animal body plan has been highly conserved throughout our evolutionary history. Biologists use similarities and differences in morphology (structure) and in early development to infer evolutionary relationships among animal groups. Variations in major characters of the body plan provide clues to animal relationships. Biologists compare variations in body symmetry, number of tissue layers, type of body cavity, and pattern of development. In addition to these traditional methods, biologists now have molecular tools to enhance our understanding of animal phylogeny.

Animals exhibit two main types of body symmetry Symmetry refers to the arrangement of body structures in relation to the body axis. Most sponges are not symmetrical, so when a sponge is cut in half, the two halves are not similar to each other. Most other animals exhibit either radial or bilateral body symmetry. Cnidarians (jellyfish, sea anemones, and their relatives) and adult echinoderms (sea stars and their relatives) have radial symmetry. In radial symmetry, the body has the general form of a wheel or cylinder, and similar structures are regularly arranged as spokes from a central axis (FIG. 30-3a). Multiple planes can be drawn through the central axis, each dividing the organism into two mirror images. An animal with radial symmetry receives stimuli equally from all directions in the environment. Some animals have modified radial symmetry. For example, sea anemones and ctenophores (comb jellies) have biradial symmetry, in which parts of the body have become specialized so that only two planes can divide the body into similar halves. Most animals exhibit bilateral symmetry, at least in their larval stages. A bilaterally symmetrical animal can be divided through only one plane (which passes through the midline of the body) to produce roughly equivalent right and left halves that are mirror images (FIG. 30-3b). As bilateral symmetry evolved, a corresponding evolutionary trend led toward cephalization, the development of a head where sensory structures are concentrated. In many animal groups, concentrations of nerve cells in the head form a brain, and a nerve cord extends from the brain toward the rear end of the animal. Bilateral symmetry and cephalization are adaptations for locomotion. The head end of the animal meets its environment first and is best equipped to capture food or respond to danger. Some definitions of basic terms and directions will help in locating body structures in bilaterally symmetrical animals. The back surface of an animal is its dorsal surface; the underside (belly) is its ventral surface. Anterior (or cephalic) means toward the head end of the animal; posterior, or caudal, means toward the tail end. A structure is said to be medial if it is located toward the midline of the body and lateral if it is toward one side of the body; for example, the human ear is lateral to the nose. In human anatomy, the term superior refers to a structure located above some point of reference, or toward the head end of the body. The term inferior

Top view

Side view

(a) Radial symmetry. Multiple planes can be drawn through the central axis; each divides the animal into two mirror images.

Dorsal

Dorsal

Frontal section

Sagittal section

Caudal

Medial Posterior

Anterior

Frontal section Lateral

Cephalic

Ventral

Cross (or transverse) section

Lateral view

Ventral

Front view

(b) Bilateral symmetry. The head of the animal is its anterior end, and the opposite end is its posterior end. The back of the animal is its dorsal surface, and the belly is its ventral surface. The diagrams illustrate various ways the body can be sectioned (cut) to study its internal structure. A sagittal section (lengthwise vertical cut) divides the animal into right and left parts. A frontal, or longitudinal, cut (lengthwise horizontal) divides the body into dorsal and ventral parts.

FIGURE 30-3 Animated Radial and bilateral symmetry Sections of animals are used in illustrations throughout this book to show the structure and arrangement of tissues and organs.

is used in human anatomy to mean located below some point of reference, or toward the feet. A bilaterally symmetrical animal has three axes, each at right angles to the other two: an anterior-posterior axis extending from head to tail, a dorsal-ventral axis extending from back to belly, and a left-right axis extending from side to side. We can distinguish three planes or sections that divide the body into specific parts. A sagittal plane divides the body into right and left parts. A sagittal plane passes from anterior to posterior and from dorsal to ventral. A frontal plane divides a bilateral body into dorsal and ventral parts. A transverse section, or cross

section, cuts at right angles to the body axis and separates anterior and posterior parts.

Animal body plans are linked to the level of tissue development Sponges have several types of cells, but they are not organized into tissues, groups of closely associated, similar cells that work together to carry out specific functions. In the early development of all animals except sponges, cells form layers, called germ layers. The outer germ layer, or ectoderm, gives rise to the tissues

that form the outer covering of the body and to nervous tissue. The inner layer, or endoderm, forms the lining of the digestive tube and other digestive structures. These layers develop into specific types of tissues. Biologists describe cnidarians and ctenophores as diploblastic because they have two tissue layers. Other animals are triploblastic. They have a third germ layer, the mesoderm, which gives rise to most other body structures, including muscles, skeletal structures, and circulatory system (when present).

Most bilateral animals have a body cavity lined with mesoderm

Epidermis (from ectoderm) Muscle layer (from mesoderm) Mesenchyme (gelatin-like tissue) Epithelium (from endoderm)

(a) Acoelomate—flatworm (liver fluke). An acoelomate animal has no body cavity. Pseudocoelom Epidermis (from ectoderm) Muscle layer (from mesoderm)

Epithelium The vast majority of bilateral animals (from endoderm) have a fluid-filled body cavity, or coelom (pronounced “see¿-lum”), between the outer wall of the body and the digestive (b) Pseudocoelomate—nematode. A pseudocoelomate animal has a body cavity that is not tube (FIG. 30-4). The flatworms and ribcompletely lined with mesoderm. bon worms are exceptions. They are bilateral and triploblastic but have a solid body; that is, they have no body cavity. True coelom Epidermis They are referred to as acoelomates (a-, (from ectoderm) “without”; and coelom, “cavity”). Muscle layer Most animals have a body cavity (from mesoderm) that is completely lined with mesoderm. Peritoneum Such a body cavity is a true coelom. An (from mesoderm) animal with a true coelom is referred to Epithelium as a true coelomate animal. The coelom (from endoderm) was one of the most important early animal adaptations. During the Proterozoic Mesentery (from mesoderm) eon, animals were small, simple forms. Evolution of the coelom was an important step in the evolution of larger, more (c) True coelomate—vertebrate. A true coelomate animal has a true coelom, a body cavity complex animals. completely lined with tissue that develops from mesoderm. With the evolution of the coelom came a new body design, the tubeFIGURE 30-4 Animated Three basic body plans in triploblastic animals within-a-tube body plan. The coelom The germ layer from which each tissue was derived is indicated in parentheses. Ectoderm is shown in blue, mesoderm in red, and endoderm in yellow. is a space that separates the body wall, the outer tube, from the digestive tube provided the opportunity for animals to become specialized in (gut), which is the inner tube. The digestive tube is attached to the swimming, crawling, or walking. Animals that can move quickly to body wall at its ends. Typically, the digestive tube has a mouth at capture food or avoid predators are more likely to survive. one end for taking in food and an anus at the other end for elimiEvolution of the coelom provided a space where internal ornating wastes. Because the coelom separates the muscles of the gans could develop and function. For example, the pumping acbody wall from those in the wall of the digestive tract, the digestive tion of the heart is possible because of the surrounding space the tube can move food along independently of body movements. coelom provides. The fluid-filled coelom also protects internal orBecause it is an enclosed compartment (or series of compartgans by cushioning them. ments) of fluid under pressure, the coelom can serve as a hydroSome (typically, small) animals have a body cavity that is static skeleton in which contracting muscles push against a tube not completely lined with mesoderm. This type of body cavity is of fluid. The hydrostatic skeleton also shapes the body of soft anicalled a pseudocoelom (“false coelom”). Animals with a pseudomals. The evolution of various shapes and divisions of the coelom

coelom, such as nematodes (roundworms) and rotifers, are called pseudocoelomates. Animals with a pseudocoelom are not a monophyletic group. They probably evolved through a process of simplification from more than one group of animals with a true coelom.

Polar axis

Top view

Bilateral animals form two main clades based on differences in development Embryonic development begins as the zygote undergoes cleavage, the first several cell divisions of the embryo. During cleavage, the embryo develops into a hollow ball of cells, the blastula. Cells of the blastula undergo gastrulation, a process that forms and segregates the three germ layers. Basic differences in the pattern of early development distinguish two main evolutionary lines of bilateral animals: protostomes (pro¿-tuh-stomes), assigned to the clade Protostomia, and deuterostomes (doo¿-ter-uh-stomes), assigned to the clade Deuterostomia. The protostomes include mollusks (for example, snails, clams, squids), annelids (for example, earthworms), arthropods (for example, crabs, insects), and several other groups. Deuterostomes include the echinoderms (such as sea stars and sea urchins) and chordates (which include the vertebrates). One important difference in the development of protostomes and deuterostomes is the pattern of cleavage. In many protostomes, the early cell divisions are diagonal to the polar axis (the long axis of the egg), resulting in a somewhat spiral arrangement of cells; any one cell lies between the two cells above or below it (FIG. 30-5a). This pattern of division is known as spiral cleavage. In radial cleavage, characteristic of the deuterostomes, the early divisions are either parallel or at right angles to the polar axis. The resulting cells lie directly above or below one another (FIG. 30-5b). In the protostomes, the developmental fate of each embryonic cell is typically fixed very early. For example, if the first four cells of an annelid embryo are separated, each cell develops into only a fixed quarter of the larva; this pattern of cleavage is called determinate cleavage. In contrast, deuterostomes typically undergo indeterminate cleavage. For example, if the first four cells of a sea star embryo are separated, each cell can form a complete, though small, larva. If a few cells are removed from a blastula undergoing indeterminate cleavage, other cells compensate, and the embryo develops normally. In contrast, if a few cells are removed from the blastula of an embryo undergoing determinate cleavage, some structure, such as a limb, does not develop. During gastrulation, a group of cells moves inward, forming a sac that becomes the embryonic gut. The opening to the outside is called the blastopore. In most protostomes, the blastopore develops into the mouth. The word protostome comes from Greek words meaning “first” and “the mouth.” In deuterostomes the blastopore does not give rise to the mouth but generally develops into the anus. A second opening that forms later in development gives rise to the mouth. The word deuterostome derives from words meaning “second” and “the mouth.” Another, though less reliable, difference between protostome and deuterostome development is the manner in which the coelom forms. In most protostomes, the mesoderm splits, and the split widens into a cavity that becomes the coelom (FIG. 30-6). This method

(a) Spiral cleavage is characteristic of protostomes. Note the spiral arrangement, with the upper cells centered between the lower cells.

Polar axis

Top view

(b) Radial cleavage is characteristic of deuterostomes. The early divisions are either parallel to the polar axis or at right angles to it. The cells are stacked, with the upper cells centered directly above the lower cells.

FIGURE 30-5 Spiral and radial cleavage The pattern of cleavage can be appreciated by comparing the positions of the purple cells in (a) and (b).

of coelom formation is known as schizocoely. In deuterostomes, the mesoderm forms as “outpocketings” of the developing gut, a process called enterocoely. These outpocketings eventually pinch off and form pouches; the cavity within the pouches becomes the coelom.

Biologists have identified major animal clades based on structure, development, and molecular data Biologists have long inferred evolutionary relationships among animals, based on the structural variations and patterns of development we have just discussed. During the past few decades, the development of sophisticated new technology, such as new types of microscopy and cell-labeling, has helped systematists in their work. Techniques for determining the sequence of nucleotides in DNA and RNA and other molecular tools have helped systematists clarify the evolutionary relationships among animal groups. Molecular data have also challenged some traditional conclusions, for example, when various animal groups diverged in relation to other groups. Such questions continue to be the topic of much debate. Animals are referred to as metazoa. Animals with two or three germ layers (diploblastic or triploblastic) make up the eumetazoa. As we have described, some animals exhibit asymmetry (sponges), radial symmetry (cnidarians), or biradial symmetry (ctenophores). The Bilateria, the lineage of animals with bilateral symmetry, are

Enterocoely— characteristic of deuterostomes

Ectoderm

Ectoderm

Developing mesoderm

Presumptive mesoderm

Blastopore

Gut

Endoderm

Enterocoelic pouch

Mesoderm

Ectoderm Endoderm

Ectoderm Ectoderm

Developing coelom (Schizocoel)

Endoderm Gut

Mesoderm Coelom (Enterocoel)

Gut

Coelom

Gut

Mesoderm

Endoderm

Mesentery

Coelom

Epidermis (ectoderm) Peritoneum (mesoderm)

Muscle layer (mesoderm) Gut

triploblastic. (They have three germ layers: ectoderm, endoderm, and mesoderm.) As explained in the last section, biologists classify bilateral animals into protostome and deuterostome clades. Molecular data show that the protostomes split into two major clades: Lophotrochozoa (pronounced “lo-fah-tro-kah-zo¿-ah”) and Ecdysozoa (pronounced “ek-dah-so-zo¿-ah”) (FIG. 30-7 and TABLE 30-1). Animals assigned to the Lophotrochozoa are characterized by (1) a lophophore, a ciliated ring of tentacles surrounding the mouth that serves as a feeding organ, or (2) a type of larva called a trochophore larva. The Lophotrochozoa include the flatworms, ribbon worms, mollusks, annelids, and three groups sometimes referred to as the lophophorate phyla.

▲

Schizocoely— characteristic of protostomes

FIGURE 30-6 Two types of coelom formation

The coelem originates in the embryo from blocks of mesoderm that split off from each side of the embryonic gut. In protostomes, the coelom typically forms by the process of schizocoely. Solid blocks of mesoderm form around the gut. Programmed cell death occurs, causing the mesoderm (red) to split. The split widens, forming a cavity that becomes the coelom. In enterocoely, characteristic of deuterostomes, the mesoderm outpockets from the gut, forming pouches. The cavity within these pouches becomes the coelom. Ectoderm is shown in blue, endoderm in yellow. The diagrams in the top row are longitudinal sections of developing embryos; other diagrams are cross sections.

The name Ecdysozoa is derived from the fact that animals in this group molt, a process called ecdysis. The Ecdysozoa include the nematodes and arthropods. Note that the phylogeny we have described assigns bilateral animals to three major clades: Lophotrochozoa, Ecdysozoa, and Deuterostomia. Segmentation apparently evolved three times Over millions of years, evolutionary forces acting on the basic animal body plan have produced changes resulting in a remarkable diversity of body forms. One very important innovation has been segmentation, a body plan in which certain structures are repeated, producing a series of body compartments. Each compartment can be regulated somewhat independently of the others, which means that various parts of the body can become specialized to perform various functions. Perhaps the most obvious example of segmentation can be found in the earthworm. However, as will be discussed in the following chapters, arthropods and vertebrates also have segmented body plans. Thus, segmented animals are found within each of the three major clades of bilateral animals. Molecular data suggest that segmentation evolved independently three times. This view is reflected in the cladogram in Figure 30-7. In each independent origin of segmentation, natural selection apparently acted on many of the same genes (for example, Hox genes). In this chapter, we have briefly discussed characteristics common to animals and the early evolution of animals. We described animal body plans and examined some of the criteria that biologists use to reconstruct phylogenetic relationships. The cladogram shown in Figure 30-7 depicts some of the current hypotheses regarding the relationships among major animals groups, and Table 30-1 summarizes these relationships. In the next two chapters, we survey these animal groups. In Chapter 31, we describe the sponges, cnidarians, ctenophores, and the protostomes. Then, in Chapter 32, we discuss the deuterostomes, which include the echinoderms and chordates.

Review ■ ■ ■

How are animals classified based on type of symmetry? What are some differences between protostomes and deuterostomes? What are the three main clades of bilateral animals, and how do they differ from one another?

KEY POINT

Three major clades of bilateral animals are recognized: Lophotrochozoa, Ecdysozoa, and Deuterostomia.

Eumetazoa Bilateria Protostomia

Deuterostomia

Chordates

Hemichordates

Echinoderms

Arthropods

Tardigrades

Nematodes

Rotifers

Lophophorates

Annelids

Mollusks

Nemerteans

Flatworms

Ctenophores

Cnidarians

Sponges

Choanoflagellates (outgroup)

Onychophorans

Ecdysozoa

Lophotrochozoa

Segmentation Segmentation

? ? Segmentation Molting

Deuterostome pattern of development

Protostome pattern of development Radial symmetry

Bilateral symmetry, three tissue layers, body cavity

?

Tissues

Deuterostomia

Ecdysozoa

Ctenophores

Cnidarians

(a) Using the criteria indicated by the colored bars above the cladogram, identify which animal groups are assigned to each clade. The evolution of some key characters is indicated on the branches. Note that segmentation evolved independently in each of the three major clades of bilateral animals.

Sponges

Choanoflagellate ancestor

Lophotrochozoa

Multicellularity

?

Choanoflagellate ancestor

(b) Six major groups of animals.

FIGURE 30-7 Evolutionary relationships of animal clades This cladogram is based on many forms of data, including comparisons of DNA sequences of several genes. Remember that any cladogram of animal relationships is a work in progress. As new data are considered, our understanding of relationships among animal groups changes.

Labels indicate when certain key traits originated. Note that the simplified cladogram shown in (b) is modified and used as an icon for many figures in Chapters 31 and 32.

TABLE 30-1

Overview of the Animal Kingdom

Major Groups and Subgroups

Some Key Characters/Comments

Poriferans (sponges)

Collar cells, flagellated cells that trap food; cells loosely associated and do not form true tissues; sponge larvae have flagella and can swim about; poriferans are probably not a monophyletic group

Cnidarians Hydrozoans: hydras Scyphozoans: jellyfish Cubozoans: box jellyfish Anthozoans: corals, sea anemones

Radial symmetry; cnidocytes (stinging cells); polyp and medusa body forms; tentacles surround mouth; mainly marine

Ctenophores (comb jellies)

Biradial symmetry; eight rows of cilia that resemble combs; tentacles with adhesive glue cells; marine predators

Protostomes: Lophotrochozoan Branch

Bilateral symmetry; triploblastic

Flatworms (platyhelminths) Turbellarians: planarians Trematodes and monogeneans: flukes Cestodes: tapeworms

Gastrovascular cavity with one opening; no coelom; cephalization

Nemerteans (ribbon worms)

Proboscis (long, muscular tube that can be everted to capture prey); complete digestive tube; mainly marine carnivores

Mollusks Chitons Gastropods: snails, slugs, nudibranchs Bivalves: clams, oysters Cephalopods: squids, octopods

Soft body usually covered by dorsal shell; muscular foot; foot; mantle covers visceral mass; most have radula (belt of teeth)

Annelids Polychaetes: sandworms, tubeworms Oligochaetes: earthworms, freshwater worms Hirudinids: leeches

Segmented body; most have bristles called setae, that provide traction during crawling

Lophophorates Brachiopods Phoronids Bryozoans (ectoprocts)

Lophophore (ring of ciliated tentacles around mouth); mainly sessile marine organisms

Rotifera (wheel animals)

Crown of cilia at anterior end; microscopic aquatic animals

Protostomes: Ecdysozoan Branch

Protostomes with cuticle that is molted and replaced as animal grows

Nematodes (roundworms)

Fluid-filled pseudocoelom serves as hydrostatic skeleton; important decomposers; many are predators

Onycophorans (velvet worms)

Sister group of arthropods? Unjointed, paired appendages

Tardigrades (“water bears”)

Sister group of arthropods? Unjointed, clawed legs

Arthropods Myriapods (centipedes, millipedes) Chelicerates (horseshoe crabs, arachnids) Crustaceans (lobsters, crabs, barnacles, copepods) Hexapods (insects)

Segmented; exoskeleton of chitin; paired, jointed appendages; insects and many crustaceans have compound eyes (continued)

TABLE 30-1

Overview of the Animal Kingdom (continued)

Major Groups and Subgroups

Some Key Characters/Comments

Deuterostomes

Radial, indeterminate cleavage; pharyngeal slits

Echinoderms Crinoids (sea lilies, feather stars) Asteroids (sea stars) Ophiuroids (basket stars, brittle stars) Echinoids (sea urchins, sand dollars) Holothuroids (sea cucumbers)

Water vascular system; tube feet; endoskeleton with spines; larvae bilateral, ciliated; adult, pentaradial symmetry; marine

Hemichordates (acorn worms)

Proboscis, collar and trunk

Chordates

Notochord; dorsal, tubular nerve cord; postanal tail; endostyle; segmented body

■ ■

Urochordates (tunicates)

Larvae have chordate characters

Cephalochordates (lancelets)

Notochord extends from anterior to posterior tip

Vertebrates (hagfishes, lampreys, cartilaginous fishes, ray-finned fishes, coelacanths, lungfishes, amphibians, reptiles [including birds], mammals)

Vertebral column, cranium, neural crest cells; endoskeleton

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

30.1 (page 627)

■

1 Describe several characteristics common to most animals. ■

■

■

Members of kingdom Animalia are eukaryotic, multicellular, heterotrophic organisms with cells specialized to perform specific functions. Animals have diverse body plans. The body plan is the basic structure and functional design of the body. Most animals are capable of locomotion at some time during their life cycle, can respond adaptively to external stimuli, and can reproduce sexually. In sexual reproduction, sperm and egg unite to form a zygote. The zygote undergoes cleavage, a series of cell divisions that produce a hollow ball of cells called a blastula. Most animals develop into a larva, a sexually immature form that may appear and behave differently from the adult. Larvae typically go through metamorphosis, a developmental process that converts the immature animal into a juvenile form that grows into an adult. Explore the characteristics of animals by clicking on the figures in CengageNOW.

30.2 (page 627) 2 Compare the advantages and disadvantages of life in the ocean, in fresh water, and on land. Marine environments have relatively stable temperatures, provide buoyancy, and provide readily available food. Fluid and salt balance are more easily maintained than in fresh water. Currents and other water movements are a disadvantage. Fresh water offers a less constant environment and less food. ■ Because fresh water is hypotonic to tissue fluid, animals must osmoregulate. ■

Terrestrial animals must have adaptations that protect them from drying out and from temperature changes, and that protect their gametes and embryos.

30.3 (page 629) 3 Use current hypotheses to trace the early evolution of animals. ■

■

Based on molecular data, biologists hypothesize that most animal clades actually diverged over a long period during the Proterozoic eon. During the Cambrian radiation, new animal body plans rapidly evolved among clades that already existed. Hox genes control early development in animal groups. These genes had evolved by the beginning of the Cambrian period, and mutations in these genes could have resulted in rapid changes in animal body plans.

30.4 (page 630) 4 Describe how biologists use morphology (including variations in body symmetry, number of tissue layers, and type of body cavity) and patterns of early development to infer relationships among animal phyla. ■ Biologists hypothesize that cnidarians (which have radial symmetry) and ctenophores (which have biradial symmetry) are more closely related to each other than to animals that exhibit bilateral symmetry. Cephalization, the development of a head, evolved along with bilateral symmetry. ■ Biologists have also inferred relationships based on level of tissue development and type of body cavity. Embryonic tissues, called germ layers, include the outer layer, ectoderm, which gives rise to the body covering and the nervous system; the inner layer, endoderm, which lines the gut and other digestive organs; and a middle layer, mesoderm, which gives rise to muscle, skeletal structures, and most other body structures.

■

In bilateral animals, the type of body cavity has been used to classify animals. Acoelomate animals have no body cavity; coelomate animals have a true coelom, a body cavity completely lined with mesoderm. Some animals have a pseudocoelom (literally, a “false cavity”), a body cavity that is not completely lined with mesoderm. Pseudocoelom

■

■

True coelom

Two major evolutionary branches of bilateral animals are Protostomia (mollusks, annelids, and arthropods) and Deuterostomia (echinoderms and chordates). Protostomes undergo spiral cleavage, in which early cell divisions are diagonal to the polar axis. Deuterostomes undergo radial cleavage, in which the early cell divisions are either parallel or at right angles to the polar axis, so the cells lie directly above or below one another.

Protostomes undergo determinate cleavage, in which the fate of each embryonic cell is fixed very early. Deuterostomes undergo indeterminate cleavage, in which the fate of each cell in early development is more flexible. ■ In protostomes the blastopore, the opening from the embryonic gut to the outside, develops into the mouth; in deuterostomes the blastopore typically becomes the anus. 5 Cite specific examples of how data from molecular systematics have confirmed or modified traditional animal phylogeny. (Include identification of three major clades of bilateral animals.) ■ Molecular systematics has confirmed much of animal phylogeny based on structural characters, including the axiom that animal body plans usually evolved from simple to complex. However, molecular systematics has also provided evidence for exceptions. ■ Based on molecular data, biologists now subdivide the protostomes into two clades: Lophotrochozoa and Ecdysozoa. The Lophotrochozoa include the flatworms, ribbon worms, mollusks, annelids, rotifers, and animals that have a lophophore, a ciliated ring of tentacles surrounding the mouth. The Ecdysozoa, animals that molt, include the nematodes and arthropods. Thus, the three major clades of bilateral animals are Lophotrochozoa, Ecdysozoa, and Deuterostomia. ■

Eumetazoa Bilateria Protostomia Lophotrochozoa

Deuterostomia Ecdysozoa

T E S T YO U R U N D E R S TA N D I N G 1. Which of the following is not a defining character of animals? (a) heterotrophic (b) multicellular (c) eukaryotic (d) presence of a coelom (e) formation of a zygote that undergoes cleavage 2. Which of the following is not an adaptation to terrestrial living? (a) internal fertilization (b) shell surrounding egg (c) adaptations for maintaining body temperature (d) surface for gas exchange deep in body (e) ability to maintain location 3. The Cambrian radiation (a) occurred during the late Cambrian period (b) was a rapid evolution of new animal body plans during the middle Cambrian (c) was a result of prokaryotic and eukaryotic migration to many new regions (d) is supported by the great variety of Ediacaran fossils (e) b and c 4. Cephalization (a) evolved along with bilateral symmetry (b) is the development of a digestive system (c) is characteristic of protostomes but not deuterostomes (d) involves a concentration of endocrine organs (e) first evolved in deuterostomes

9. The evolution of animals (a) followed an orderly progression from simple to complex (b) will be better understood as biologists continue to collect molecular and other types of data (c) has been determined by studying classification (d) began with the cnidarians (e) began with their common ancestor, a protostome 10. Which of the following would be most helpful in deciding whether an animals is a lophotrochozoan? (a) undergoes metamorphosis (b) molts (c) its zygote undergoes a series of mitotic divisions and becomes a blastula (d) has a trochophore larva (e) forms three germ layers 11. Which of the following is an example of a deuterostome? (a) lophotrochozoan (b) coral (c) chordate (d) planarian flatworm (e) insect 12. Label the branches of the diagram. Use Figure 30-7b to check your answers.

5. The germ layer that gives rise to the outer covering of the body and the nervous system is the (a) gastrodermis (b) ectoderm (c) mesenchyme (d) endoderm (e) mesoderm 6. Radial symmetry is characteristic of (a) protostomes (b) chordates (c) deuterostomes (d) cnidarians (e) poriferans 7. A true coelom is completely lined with (a) mesenchyme (b) ectoderm (c) mesoderm (d) endoderm (e) epidermis 8. Protostomes are characterized by (a) spiral cleavage (b) indeterminate cleavage (c) enterocoely (d) radial symmetry (e) a and c

?

CRITICAL THINKING 1. Imagine that you discover a new organism in a lake. How would you decide whether it is an animal? What are some characteristics that might contribute to your decision? 2. EVOLUTION LINK. Examine the cladogram in Figure 30-7. Based on the discussion in this chapter, what were some of the types of data that biologists used to determine these phylogenetic relationships? 3. EVOLUTION LINK. Suggest and support a hypothesis for the view that segmentation evolved in three different animal clades.

4. ANALYZING DATA. Imagine that a biologist discovers evidence that a type of sponge evolved from an early protostome. Where would you place the branch for that sponge on Figure 30-7a? Additional questions are available in CengageNOW at www.cengage.com/ login.

31

Sponges, Cnidarians, Ctenophores, and Protostomes

Marty Snyderman/Visuals Unlimited

A bearded fireworm (Hermodice carunculata). The fireworm is a segmented worm (an annelid) that can grow up to 31 cm (11.8 in) long. It lives in shallow marine waters—in coral reefs, in beds of turtle grass, or under rocks. Fireworm bristles, which are filled with venom, can break off in the skin and cause irritation.

KEY CONCEPTS

I

n Chapter 30 we introduced the animal kingdom and explored animal phylogeny. We examined how systematists use fossils, morphology,

31.1 Animals with asymmetry, radial, or biradial symmetry

developmental patterns, and molecular data to determine animal rela-

include sponges, cnidarians (hydras, jellyfish, sea anemones), and ctenophores (comb jellies). Sponges are characterized by collar cells and by loosely associated cells that do not form true tissues; cnidarians are characterized by radial symmetry, two tissue layers, and cells that contain stinging organelles; and ctenophores have biradial symmetry, two tissue layers, eight rows of cilia, and tentacles with adhesive glue cells.

tionships. In this chapter we begin with an introduction to the sponges,

31.2 Protostomes are a monophyletic group that gave rise to two major clades: Lophotrochozoa and Ecdysozoa. The lophotrochozoans include the annelids (earthworms), mollusks, and several smaller groups.

31.3 The ecdysozoans include the nematodes (roundworms) and arthropods (horseshoe crabs, spiders, insects). The remarkable biological success of the insects can be attributed to the evolution of complex body plans and life cycles including their exoskeleton, segmentation, specialized jointed appendages, ability to fly, and metamorphosis.

cnidarians, and ctenophores, groups characterized by asymmetry, radial symmetry, and biradial symmetry, respectively. The phylogeny of these animals is the focus of much current research. Although sponges, cnidarians, and ctenophores are mainly small animals with simple body structures, they are of great ecological importance. They are important members of marine food chains, many provide shelter for other organisms, and some form symbiotic relationships with other animals. Corals that produce reefs are among the most ecologically important animals in the world. Following our discussion of the sponges, cnidarians, and ctenophores, we begin our survey of the bilateral animals, the most familiar and by far the most numerous animals on Earth. More than 99% of animal species belong to the Bilateria. These diverse forms have adapted to life in almost every imaginable habitat in the ocean and in freshwater and terrestrial

environments. The bearded fireworm shown in the photograph is an example of the diverse animals found among the Bilateria. The remarkable success of the bilateral animals can be attributed to the evolution of adaptations that facilitated food capture, escape from predators, and reproduction. Among their key adaptations are cephalization (formation of a head), a central nervous system, muscles, a coelom, and compartmentalization of the body. Large animals developed body systems for gas exchange, waste disposal, and internal transport of nutrients, respiratory gases, wastes, and other materials. Recall that based on pattern of early development, bilateral animals are classified as protostomes or deuterostomes. The protostomes form two clades: Lophotrochozoa and Ecdysozoa. In this chapter, we focus on the innovations in body plans, phylogenetic relationships, and life histories of several phyla of these two clades. For reasons discussed in Chapter 23, some systematists now avoid using fixed ranks of the Linnaean classification hierarchy (kingdom, phylum, class, order, family). In classifying animals, we describe clades when these are known. We continue to use some Linnaean ranks, noting groups that may not be monophyletic and pointing out the controversies.

31.1 SPONGES, CNIDARIANS, AND CTENOPHORES: ANIMALS WITH ASYMMETRY, RADIAL, OR BIRADIAL SYMMETRY ■ ■ LEARNING OBJECTIVES 1 2 3

Identify distinguishing characteristics of poriferans. Identify distinguishing characteristics of cnidarians, describe four groups, and give examples of animals that belong to each group. Identify distinguishing characteristics of ctenophores.

Until recently, biologists hypothesized that sponges diverged early from other animal groups and they assigned sponges to a basal group, Parazoa (para, “alongside”; and zoa, “animals”). They based their hypothesis on the asymmetry and simple body plan of sponges. The cells of sponges are loosely associated and do not form true tissues. Other animals have true tissues and were classified as Eumetazoa (eu, “true”; meta, “later”; and zoa, “animals”). Thus, biologists divided the animal kingdom into two major animal clades: Parazoa and Eumetazoa. Parazoa was considered a sister group of the Eumetazoa; however, this view has been challenged. At least one group of sponges (calcareous sponges) may have become simplified as they evolved from animals with more complex body plans. Some systematists have proposed that comb jellies, rather than sponges, may be the oldest group of animals. Systematists continue to search for new clues that will settle the controversies over which branch of animals is the oldest, as well as which groups are more closely related to one another.

Sponges have collar cells and other specialized cells Molecular data suggest that the poriferans (paw-rif ¿-er-ans; phylum Porifera) are not a monophyletic group. The name Porifera, meaning “to have pores,” aptly describes the sponges, whose bodies are perforated by tiny holes. Sponges are aquatic, mainly marine animals that are most abundant in warm waters. Biologists have identified about 15,000 species of sponges. They range in size from a few millimeters to more than a meter in height and diameter (loggerhead sponges). Many sponges are asymmetrical, but they vary in shape from flat, encrusting growths to balls, cups, fans, or vases. Living sponges may be brightly colored—green, orange, red, yellow, blue, or purple—or they may be white or drab (FIG. 31-1). Some species are inhabited by symbiotic bacteria or algae that give them color. Although they are multicellular and can be large, sponges function much like choanoflagellates—a group of unicellular and colonial protists. Recall that evidence suggests that animals (including sponges) and choanoflagellates share a common choanoflagellate ancestor. Choanoflagellates are characterized by a single flagellum surrounded by a collar of microvilli (see Fig. 26-21). Sponges have flagellate cells called collar cells, or choanocytes, which are strikingly similar to choanoflagellates. Sponge larvae have flagella and can swim about. Adult sponges attach to some solid object and have long been described as sessile. However, biologists have observed adults of several species moving slowly (about 4 mm per day), perhaps by the cumulative movement of cells along the sponge’s lower surface. Although sponges are multicellular, their cells are loosely associated and do not form true tissues. However, a division of labor exists among the several types of cells that make up the sponge, with certain cells specializing in nutrition, support, contraction, or reproduction. Many sponge cells are extremely versatile and can change form and function. Collar cells make up the inner layer of certain sponges. Each cell is equipped with a tiny collar surrounding the base of the flagellum. The collar is an extension of the plasma membrane and consists of microvilli. Collar cells create the water current that brings food and oxygen to the cells and carries away carbon dioxide and other wastes. Collar cells also trap and phagocytize food particles. Together, the collar cells of some sponges can pump a volume of water equal to the volume of the sponge each minute! Sponges have three types of canal systems through which water circulates. In the simple asconoid canal system, the beating flagella of collar cells create a current that pulls water through hundreds of tiny pores, called ostia. Specialized tubelike cells, called porocytes, form the pores. These cells regulate the diameter of the pores by contracting. Water passes into the central cavity, or spongocoel (not a digestive cavity), and then flows out through the sponge’s open end, the osculum. The asconoid system is illustrated in Figure 31-1. In the syconoid system, water enters through canals lined by collar cells. Most types of sponges have a leuconoid system. The body wall is extensively folded, and complex systems of canals provide increased surface area for food capture. Epidermal cells form the outer layer of the sponge and line the canals. The canals lead into small chambers that are lined with collar cells.

Deuterostomia

Ecdysozoans

Lophotrochozoans

Ctenophores

Sponges

Cnidarians

Osculum

Water movement

Spongocoel ?

Choanoflagellate ancestor

Ostia (incurrent pores)

Charles Seaborn / Odyssey Productions, Chicago

Epidermal cell

(b) Porocyte

Spicule Microvillus

(a)

Nucleus

Flagellum

Collar cell (choanocyte)

Amoeboid cell in mesohyl

Collar

FIGURE 31-1 Animated Sponge structure (a) Tube sponges (Spinosella plicifera) from the Caribbean, attached to the coral reef substrate. (b) Sponge cut to expose its organization. This sponge has an asconoid type of canal system.

The skeletal framework of a sponge can be both rigid and fibrous. Between the outer and inner cell layers, the sponge body has a gelatin-like layer, the mesohyl, which is supported by slender skeletal spikes, or spicules. Amoeboid cells, which wander about in the mesohyl, secrete the spicules. The spicules are made of calcium carbonate or of silica. The fibrous part of the sponge skeleton consists of spongin, a form of collagen. Sponges are suspension feeders, adapted for trapping and eating whatever food is suspended in the water. As water circulates through the body, food is trapped along the sticky collars of the collar cells. Thus, the collars filter the suspended food particles out of the water. Suspension feeders that filter the suspended food are known as filter feeders. Food particles are either digested within the collar cell or transferred to an amoeboid cell for digestion. The amoeboid cell transports nutrients to epidermal cells. Undigested food passes out through the osculum and is simply eliminated into the water. Gas exchange and excretion of wastes depend on diffusion into and out of individual cells. Although cells of the sponge can

react to stimuli, sponges do not have specialized nerve cells and so cannot react as a whole. However, electrical signals have been identified in certain glass sponges. Behavior appears limited to basic metabolic necessities such as capturing food and regulating the flow of water through the body. Sponges reproduce both asexually and sexually. In asexual reproduction, a small fragment or bud may break free from the parent sponge and give rise to a new sponge. Such fragments may attach to the parent sponge, forming, or becoming part of, a colony. Most sponges are hermaphrodites, meaning the same individual can produce both eggs and sperm. Some of the amoeboid cells develop into sperm cells; others, into egg cells. However, hermaphroditic sponges usually produce eggs and sperm at different times, and they cross-fertilize with other sponges. They release mature sperm into the water, which are taken in by other sponges of the same species. Fertilization and early development take place within the jellylike mesohyl. Zygotes develop into flagellate larvae that leave the parent along with the stream of outflowing water. After swimming

Deuterostomia

Ecdysozoans

Lophotrochozoans

Ctenophores

Cnidarians

Sponges

for a while, a larva finds a solid object, attaches to it, and settles down to a sessile life. Sponges have a remarkable ability to repair themselves when injured and to regenerate lost parts. When the cells of a sponge are separated experimentally, they recognize one another and their place in the whole and aggregate to re-form a complete sponge. There are three main groups of sponges: the calcareous sponges have spicules composed of calcium carbonate, whereas the glass sponges and the demosponges have spicules of silicon. About 95% of living sponges are demosponges. In some of the demosponges, spicules are bound together by spongin. Some demosponges do

?

Choanoflagellate ancestor

not have spicules. These soft sponges have been used for thousands of years as bath sponges and for cleaning. Unfortunately, as a result of overfishing, bath sponges are now in short supply. Molecular data indicate that the calcareous sponges are a separate clade from the glass sponges and demosponges. The calcareous sponges may be the sister taxon to the other sponges, or they may be more closely related to lophotrochozoans than to the glass sponges and demosponges.

Cnidarians have unique stinging cells About 10,000 species of cnidarians (ni-dah¿-ree-ans; phylum Cnidaria) have been described. Most are marine. The radially symmetrical cnidarian body is organized as a hollow sac with the mouth and surrounding tentacles located at one end. Some cnidarians live a solitary existence, whereas many others, such as corals, form colonies. Based on molecular data, some systematists have proposed that cnidarians share a common ancestor with the clade of bilateral animals. Cnidarians have two body shapes: polyp and medusa (FIG. 31-2). The polyp form, represented by Hydra, typically has a dorsal mouth surrounded by tentacles. In the medusa (pl., medusae), or jellyfish form, the mouth is located in the lower concave, or oral, surface; the

Mouth Mouth Mouth

Mesoglea Gastrodermis Epidermis

Epidermis

Gastrovascular cavity

Mesoglea

Mesoglea

Gastrodermis

Gastrodermis Gastrovascular cavity

(a) Class Hydrozoa (polyp). This hydrozoan (Gonothyraea loveni ) forms a colony of polyps. The drawing illustrates a single polyp.

Mike Bacon/Tom Stack & Associates

Brian Parker/Tom Stack & Associates

Robert Brons/Biological Photo Service

Gastrovascular cavity

(b) Class Scyphozoa (medusa). The sea nettle (Chrysaora fuscescens ) uses its tentacles equipped with cnidocytes to capture small animals (zooplankton) suspended in the water.

FIGURE 31-2 Animated Polyp and medusa body forms of cnidarians

(c) Class Anthozoa (polyp). Coral polyps (Montastrea cavernosa) extended for feeding.

Robert and Linda Mitchell

convex upper surface is the aboral surface. Some cnidarians have the polyp shape during one stage of their life cycle and the medusa form during another stage. The Portuguese man-of-war and some other cnidarians consist of colonies of many individuals, some of which are polyps and others medusae. Cnidarians get their name from specialized cells, called cnidocytes (from a Greek word meaning “nettle cells”), that contain stinging organelles. Cnidocytes are located mainly in the epidermis, especially on the tentacles. The cnidocytes contain stinging “thread capsules,” called nematocysts (FIG. 31-3). Each cnidocyte has a small, projecting trigger (cnidocil) on its outer surface and a coiled, hollow thread inside. We can compare the cnidocyte to a loaded gun, ready to fire. When stimulated by touch or certain chemicals

250 μm

(a) LM of discharged nematocysts of a Portuguese man-of-war (Physalia physalis ). Photographed in the Gulf of Mexico.

Cnidocyte

dissolved in the water, the nematocyst fires its thread. Some types of nematocyst threads are sticky. Others are long and coil around prey. A third type bears barbs or spines that can inject a protein toxin that paralyzes prey animals, such as small crustaceans. Cnidarians use their tentacles to capture prey and push it into the mouth. The mouth leads into the gastrovascular cavity, where digestion takes place. The mouth is the only opening into the gastrovascular cavity and so must serve for both ingestion of food and expulsion of undigested material. Gas exchange and excretion occur by diffusion. The body wall is thin enough that no cell is far from the surface. More highly organized than sponges, cnidarians are diploblastic; that is, they have two definite tissue layers. The ectoderm gives rise to the outer epidermis, a protective layer covering the body. The endoderm gives rise to the inner gastrodermis, which lines the gastrovascular cavity and functions in digestion. These thin layers are separated by a thick, jellylike mesoglea, which is mainly acellular. Cnidarians have nerve cells that form nerve nets connecting sensory cells in the body wall to contractile cells and gland cells. An impulse set up by one sensory cell passes in all directions more or less equally. Nerve cells are not organized to form a brain or nerve cord. In some types of medusae, sense organs are positioned around the edge of the body. The sense organs include simple “eyes,” called eyespots, organs of balance (statocysts), and chemoreceptors (cells that sense certain chemicals). Both the epidermis and gastrodermis have cells specialized to contract (however, they are not true muscle cells). Contractile fibers in the epidermal cells are arranged lengthwise, and those in the gastrodermis are arranged in a circular pattern. These two sets of contractile cells act on the water-filled gastrovascular cavity, forming a hydrostatic skeleton. This skeleton supports the body and allows movement. By contracting one set of contractile cells or the other, the hydra can shorten, lengthen, or bend its body (see Fig. 40-2). We consider four groups of cnidarians: hydrozoans, scyphozoans, cubozoans, and anthozoans (TABLE 31-1).

Nucleus Thread

TABLE 31-1

Capsule

Class and Representative Animals

Nematocyst (not discharged) Cnidocil (trigger)

Thread Nematocyst (discharged)

(b) Undischarged and discharged nematocyst. The cnidocil, or trigger, is a mechanoreceptor that discharges the nematocyst when it senses contact with an object.

FIGURE 31-3 Animated Nematocysts When cnidarian stinging cells (cnidocytes) are stimulated, the nematocyst discharges, ejecting a thread that may entangle or penetrate the prey. Some nematocysts secrete a toxic substance that immobilizes the prey.

Hydrozoa Hydra, Obelia, Portuguese man-of-war Scyphozoa Jellyfish Cubozoa “Box jellyfish”

Anthozoa Sea anemones, corals, sea fans

Major Classes of Phylum Cnidaria

Characteristics Mainly marine, but some freshwater species; alternation of polyp and medusa stages in most species (polyp form only in Hydra); some form colonies Mainly marine; typically inhabit coastal water, free-swimming medusa most prominent form; polyp stage often reduced Inhabit tropical and subtropical waters; have polyp stage, but medusa form most prominent; square shape when viewed from above; actively hunt prey; complex eyes that form blurred images Marine; solitary or colonial polyps; no medusa stage in most; gastrovascular cavity divided by partitions into chambers, increasing area for digestion

Biophoto Associates/Photo Researchers, Inc.

Most hydrozoans form colonies Hydrozoans include hydras and hydroids, such as Obelia and the Portuguese man-of-war Although not really typical, the solitary Hydra is the cnidarian that beginning Tentacles biology students most often study Cnidocytes (FIG. 31-4). To the naked eye, Hy(stinging cells) dra looks like a bit of frayed string. This tiny hydrozoan is found in freshwater ponds. Because it has a remarkable ability to regenerate, biologists named Hydra after the multiheaded monster of Greek Mouth mythology that could grow two 17 mm new heads for each head cut off. When Hydra is cut into several (a) Hydra viridis with large bud. When the pieces, each piece can regenerate bud separates from its parent, it becomes an independent individual. all the missing parts and become a Egg whole animal. Bud (ovum) Hydra lives in fresh water and typically attaches to a rock, aquatic plant, or detritus by a disc Gastrovascular of cells at its base. Hydras reprocavity Ovary duce asexually by budding durEpidermis ing periods when environmental Mesoglea conditions are optimal. However, Gastrodermis they differentiate as males and females and reproduce sexually in the fall or when pond water be(c) Cross section through comes stagnant. The zygote may body of Hydra. become covered with a shell that protects it through the winter or (b) Hydra cut longitudinally to show internal strucuntil conditions become more ture. Asexual reproduction by budding is represented on left; sexual reproduction is represented favorable. by ovary on right. Male hydras develop testes that Many hydrozoans form colproduce sperm. onies consisting of hundreds or thousands of individuals. A colony begins with a single polyp FIGURE 31-4 Hydra, a freshwater hydrozoan that reproduces asexually by budding. However, instead of separathave a thick, viscous mesoglea that gives firmness to the body. In ing from the parent, the bud remains attached and eventually forms scyphozoans, the polyp stage is small and inconspicuous or may additional buds. Several types of individuals may arise in the same even be absent. The largest jellyfish, Cyanea, may be more than colony, some specialized for feeding, some for reproduction, and 2 m (6.5 ft) in diameter and have tentacles 30 m (98 ft) long. These others for defense. orange and blue “monsters,” among the largest invertebrates, can Some marine hydrozoans are remarkable for their alternation cause painful stings to swimmers in the North Atlantic and Pacific of sessile polyp and motile medusa stages. The life cycle of the coOceans. lonial marine hydrozoan Obelia illustrates alternation of polyp and medusa stages (FIG. 31-5). In Obelia, polyps reproduce asexually by budding, whereas the medusae reproduce sexually. Both polyp Cubozoans include the “box jellyfish” and medusa are diploid; only sperm and eggs are haploid. When viewed from above, cubozoans have a square shape. They have four tentacles, or groups of tentacles. These fast-swimming Scyphozoans are the “true” jellyfish jellyfish have complex eyes that form blurred images, and they actively hunt for prey. Found in waters off the northern Australian In scyphozoans, the medusa is the dominant body form. Jellyfish coast, the sea wasp (Chironex fleckeri) has long tentacles that can medusae are generally larger than hydrozoan medusae, and they

extend more than 3 m (10 ft). The sea wasp produces one of the deadliest venoms in the animal kingdom. Its venom contains toxins that can stun or kill fish and other prey. These toxins can also cause respiratory failure and cardiac arrest in humans. Fatalities are mainly associated with severe stings caused by contact with tentacles over a large area of the body. Interestingly, sea turtles feed on box jellyfish, apparently unaffected by its venom.

Reproductive polyp

David M. Phillips/Visuals Unlimited

Feeding polyp

250 μm

(a) LM of Obelia. Some polyps have tentacles and are specialized for feeding, whereas others are specialized for reproduction.

1

Anthozoans are polyps Anthozoans, which include the sea anemones and corals, are either individual or colonial polyps. There is no free-swimming medusa stage. The polyp produces eggs and sperm, and the fertilized egg develops into a small, ciliated larva called a planula. This larval form may swim to a new location before attaching to develop into a polyp. Anthozoans differ from hydrozoans in that a series of vertical partitions partially divides the gastrovascular cavity into connected chambers. The partitions increase the surface area for digestion, enabling an anemone to digest an animal as large as a crab. Although corals can capture prey, many tropical species depend for nutrition on photosynthetic algae (zooxanthellae) that live within cells lining the coral’s digestive cavity (see Chapter 26 and Fig. 54-12). The relationship between coral and zooxanthellae is symbiotic and mutually beneficial. The algae provide the coral with oxygen and with carbon and nitrogen compounds. In exchange, the coral supplies the algae with waste products such as ammonia, from which the algae make nitrogenous compounds for both partners. In warm, shallow seas, much of the bottom is covered with coral or anemones, most of them brightly colored. Coral reefs con-

Reproductive polyps produce medusae by budding asexually Medusae

Mouth Tentacle Feeding polyp

2

Medusa bud Reproductive polyp

Free-swimming medusae reproduce sexually.

Sperm

Egg

Gastrovascular cavity

3

Zygote develops into ciliated planula larva.

Planula larva

Polyp colony 5

Colony grows as new polyps bud and remain attached.

4

Young polyp colony

(b) Life cycle of Obelia.

FIGURE 31-5 Animated Obelia, a marine colonial hydrozoan

Larva develops into polyp that forms new colony.

Photo by Ove Hoegh-Guldberg/Centre for Marine Studies, University of Queensland

sist of colonies of millions of corals and of certain algae (mainly coralline red algae). Living colonies occur only in the uppermost regions of such reefs, adding their own skeletons to the forming rock. Coral reefs are among the most productive of all ecosystems, rivaling tropical rain forests in species diversity (see Fig. 56-21). A single reef can serve as home for more than 3000 species of fishes and other marine organisms, and an estimated one-fourth of all marine species depend on coral reefs. A 2008 study of 704 species of reef-building corals reported that about one-third were found to be threatened or nearthreatened, and may be at increased risk for extinction. Human activities are responsible for much of this damage. Contributing factors include overfishing, mining reefs for building materials, polluting coastal waters with industrial chemicals, and smothering coral with the silt washed downstream from clear-cut forests. Acidification of the ocean caused by the increase in carbon dioxide in the atmosphere is another important factor. Coral bleaching is the stress-induced loss of the colorful symbiotic algae that inhabit coral cells (FIG. 31-6). When under environmental stress, coral actually expel the algae. Without their algae, coral become malnourished and die. Environmental factors suspected of contributing to coral bleaching include unusually low or high (associated with global warming) temperatures, pollution, changes in salinity, acidification, and disease.

Comb jellies have adhesive glue cells that trap prey

David Wrobel /Biological Photo Service

Deuterostomia

Ecdysozoans

Lophotrochozoans

Ctenophores

Sponges

Cnidarians

FIGURE 31-6 Bleached staghorn coral The fragile, luminescent comb jellies are marine animals known This staghorn coral (Acropora) was photographed off Heron Island on the as ctenophores. They are assigned to phylum Ctenophora (tehGreat Barrier Reef off the coast of Australia. nof ¿-er-uh). Approximately 150 species of ctenophores have been described. Some are as small as a pea; others are larger than Because of their similarities, biologists classify ctenophores a tomato. The outer surface of a ctenophore bears eight rows of near the cnidarians. However, their development is different, and cilia that resemble combs (FIG. 31-7). Ctenophores are biradially symmetrical, meaning you could obtain equal halves by cutting their digestive cavity has openings at both ends. The similarities through the body axis in two different ways. The ctenophore body between ctenophores and cnidarians may be a result of convergent plan is somewhat similar to that of a cnidarian medusa in that evolution from living in a similar environment, the ocean. Where, ctenophores have a type of radial symmetry, feeding tentacles, and then, do the ctenophores belong on our cladogram? As systemtwo cell layers separated by a thick, jellylike mesoglea. atists gather new data, their position on the animal tree will beSome ctenophores have two tentacles. Their tentacles are come clearer. equipped with adhesive glue cells. When prey comes in contact with the tentacle, the glue cells burst open, releasing sticky threads that trap the prey. Ctenophores do not have the stinging nematocysts characteristic of the cnidarians. Also unlike cnidarians, the ctenophore digestive system has a mouth for food intake at one ? end and two anal pores for the egestion of water and wastes at the other end. Ctenophores have a nervous system Choanoflagellate ancestor that includes a network of nerve cells and a sensory organ called a “statocyst,” which coordinates beating of the cilia in the combs. The coordinated beating of the cilia moves the animal through the FIGURE 31-7 Ctenophore (comb jelly) water. The statocyst also functions in balCtenophores are free-swimming, bioluminescent hermaphrodites capable of self-fertilization. The sea gooseberry (Pleurobrachia) has two long tentacles with adhesive cells used to capture prey. ance and helps the animal orient itself.

Review ■ ■ ■ ■ ■

Why are choanocytes significant? What is their function in sponges? In what ways do the cells of sponges specialize? In what ways do cnidarians differ from sponges? Sketch the major events of the life cycle of Obelia. How is a ctenophore like a cnidarian? In what ways is it different?

the evolution of a head with a concentration of sense organs at the anterior end. A simple brain and paired sense organs are concentrated in the “head” region. An animal with a front end generally moves forward. With a concentration of sense organs in the part of the body that first meets its environment, the animal can actively search for food, shelter, and mates or detect enemies quickly.

Flatworms are bilateral acoelomates

31.2 THE LOPHOTROCHOZOA ■ ■ LEARNING OBJECTIVES 4 Summarize what is known about the phylogeny of the lophotrocho-

5

6 7 8

zoans and cite the adaptive advantages of having a coelom and of cephalization. Identify distinguishing characteristics of flatworms and nemerteans; describe the main groups of flatworms, giving examples of animals that belong to each class. Describe the distinguishing characteristics of mollusks and of the four main groups of mollusks discussed, giving examples of each group. Describe the distinguishing characteristics of annelids and of the three main groups of annelids discussed, giving examples of each group. Describe the distinguishing characteristics of the lophophorates and of the rotifers.

We begin our discussion of the bilateral animals with the Lophotrochozoa, a clade that includes the flatworms (platyhelminths), rotifers (wheel animals), nemerteans (ribbon worms), mollusks, annelids (earthworms), and the lophophorate phyla (see Fig. 30-7). Recall from Chapter 30 that the name Lophotrochozoa comes from two important characters of some animals in this clade: (1) the lophophore, a ciliated ring of tentacles surrounding the mouth in three small groups of animals; and (2) the trochophore larva, a type of larva that characterizes two major groups—the mollusks and annelids. Some systematists argue that flatworms and rotifers should be assigned to a separate group, the platyzoa. Here, we include them with the lophotrochozoa. Most lophotrochozoans have bilateral symmetry, at least in their larval stages. They are triploblastic; that is, they have three definite tissue layers. In addition to the outer epidermis derived from ectoderm, and an inner endodermis derived from endoderm, lophotrochozoans have a middle tissue layer that develops from mesoderm. Most lophotrochozoans have a true coelom and a tube-withina-tube body plan (see Chapter 30). Most coelomate animals have well-developed circulatory, excretory, and nervous systems. Many internal organs are suspended within folds of the tissue lining the coelom and can move independently of the outer body wall. In addition, the coelom provides space for the gonads to develop. During the breeding season of many animals, such as birds, the gonads enlarge within the coelom as they fill with ripe gametes. In some animals, fluid within the coelom helps transport materials such as food, oxygen, and wastes. Cells bathed by the coelomic fluid can exchange materials with it. The cells receive nutrients and oxygen from the coelomic fluid and excrete wastes into it. Some coelomates have excretory structures that remove wastes directly from the coelomic fluid. Animals that move about typically have an elongated body with definite anterior and posterior ends. They exhibit cephalization,

The flatworms (phylum Platyhelminthes [plat-ee-hel-min¿-theez]) are soft-bodied animals that left few fossils to provide clues to their evolutionary history. Their flat, elongated bodies are solid; that is, they are acoelomate (they have no body cavity). Historically, biologists considered flatworms to be the simplest bilaterally symmetrical, triploblastic animals. However, molecular data, specifically 18S ribosomal RNA sequences, indicate that phylum Platyhelminthes, as traditionally classified, is not a monophyletic group. Based on molecular data, systematists have reclassified two orders (from class Turbellaria) that differ significantly from other flatworms. These two orders have been assigned to their own phylum (Acoelomorpha). This new group may be closely related to the common ancestor that linked radially symmetrical animals with bilateral animals. The remaining flatworms appear more closely related to coelomates. If so, flatworms evolved from more complex animals. They originally had a body cavity, but their body plan became simplified later in their evolutionary history. Many biologists currently classify these flatworms as lophotrochozoans. The phylogeny of flatworms remains controversial. As biologists consider new data, their assessment of the phylogenetic relationships of flatworms will likely change. Some biologists assign the 20,000 Platyhelminthes species to four classes. Class Turbellaria consists of the free-living flatworms, including planarians and their relatives. Classes Trematoda and Monogenea include the flukes, which are either internal or external parasites. Class Cestoda includes the tapeworms, which as adults are intestinal parasites of vertebrates (TABLE 31-2). Flatworms typically have a simple nervous system. The brain consists of two masses of nervous tissue, called ganglia, in the TABLE 31-2 Class and Representative Animals Turbellaria* Planarians Trematoda and Monogenea Flukes Cestoda Tapeworms

Classes of Phylum Platyhelminthes

Characteristics Mainly free-living; mainly marine; body covered by ciliated epidermis; typically carnivorous; prey on tiny invertebrates Parasites with a wide range of vertebrate and invertebrate hosts; may require intermediate hosts; adults have suckers for attachment to host Parasites of vertebrates; complex life cycle usually with one or two intermediate hosts; larval host may be invertebrate; typically have suckers and sometimes hooks for attachment to host; eggs produced within proglottids, which are shed; no digestive or nervous systems

*Systematists view Turbellaria as a paraphyletic group, and it will amost certainly be divided into at least three groups.

that live in digestive tracts resist the digestive enzymes secreted by their hosts. Many have complicated life cycles and produce large numbers of eggs. Other adaptations include the loss of certain structures such as sense organs. Tapeworms have also lost the digestive system.

head region. In many species, the ganglia connect to two nerve cords that extend the length of the body. This nervous system is sometimes referred to as a “ladder-type nervous system” because a series of nerves connects the cords like the rungs of a ladder (see Fig. 42-2). Sense organs include simple “eyes,” called eyespots, and organs of balance (statocysts). Flatworms have no organs for circulation or gas exchange. These functions depend largely on diffusion through the body wall. The activities of some organs are coordinated and form simple organ systems, for example, the digestive and nervous systems. As in the cnidarians, the digestive system is a gastrovascular cavity with only one opening, a mouth. The gastrovascular cavity is often extensively branched. Parasitic flatworms—flukes and tapeworms—are highly adapted to and modified for their parasitic lifestyle. They have suckers or hooks for holding on to their hosts. The bodies of those

Turbellarians are free-living flatworms Most members of class Turbellaria are free-living, marine flatworms. However, many inhabit freshwater habitats, and some are terrestrial. Turbellarian flatworms typically have a muscular pharynx that takes in food; a simple brain, eyespots, and other sensory organs in the head; protonephridia, structures that function in osmoregulation (fluid balance) and metabolic waste disposal (excretion); and reproductive organs. Planarians are turbellarian flatworms found in ponds and quiet streams throughout the world. The common American planarian Dugesia is about 15 mm (0.6 in) long, with what appear to be crossed “eyes” and flapping “ears” called auricles (FIG. 31-8). The auricles actually serve as organs of chemoreception, important in locating food. Planarians are capable of learning. Memory is not localized within the ganglia but appears to be retained throughout the nervous system. Planarians are carnivorous; they trap small animals in a mucous secretion. The digestive system consists of a single opening (the mouth); a tubelike, muscular pharynx (the first portion of the digestive tube); and a branched gastrovascular cavity. A planarian projects its pharynx outward through its mouth, using it to suck in prey. The long, highly branched gastrovascular cavity distributes food to all parts of the body so each cell can receive nutrients by diffusion. Although excretion takes place mainly by diffusion, some metabolic wastes are excreted by the protonephridia. Protonephridia

Deuterostomia

Ecdysozoa

Rotifers

Lophophorates

Annelids

Mollusks

Nemerteans

Platyhelminthes

Cniderians

Ctenophores

Sponges

Lophotrochozoa

Choanoflagellate ancestor

Ganglia

Auricle

Auricle

Pharyngeal sheath cavity Pharyngeal cavity

Eyespot Nerve Gastrovascular cavity Pharynx Sheath surrounding pharynx

Outer muscle layer of pharynx

Muscle

T. E. Adams /Peter Arnold, Inc.

Epidermis

Mouth

1 mm

(a) Internal structure.

Inner muscle layer of pharynx

(b) LM of a living planarian (Dugesia dorotocephala). Note the prominent auricles, used to locate food.

FIGURE 31-8 Animated The common planarian, Dugesia

Sperm mass

Ventral nerve cords

Adhesive gland Cilia

Body wall Muscle composed of layers epidermis, circular muscle, and longitudinal muscle

(c) Cross section through a planarian.

are blind tubules that end in flame bulbs, collecting cells equipped with cilia. The beating of the cilia channels waste through the system of tubules and eventually out of the body through excretory pores. Osmoregulation and protonephridia are discussed further in Chapter 48 (see Fig. 48-2). Planarians reproduce either asexually or sexually. In asexual reproduction, an individual constricts in the middle and divides into two planarians. Each regenerates its missing parts. Sexually, these animals are hermaphrodites. During the warm months of the year, each is equipped with a complete set of male and female organs. Two planarians come together in copulation and exchange sperm cells so that their eggs are cross-fertilized. Flukes parasitize other animals Although their body plan resembles that of the free-living flatworms, the flukes, members of classes Trematoda and Monogenea, have structures, such as hooks and suckers, for attachment to their host. Flukes also have extremely prolific reproductive organs.

Flukes that are parasitic in humans include blood flukes, widespread in tropical areas of the world, and liver flukes, common in Asia, particularly in areas where humans use their own feces for fertilizing crops. Blood flukes of the genus Schistosoma infect about 200 million people who live in tropical areas. Both blood flukes and liver flukes go through complicated life cycles involving several different forms. There is an alternation of sexual and asexual stages, and parasitism on one or more intermediate hosts, such as snails and fishes (FIG. 31-9). The aquatic snails that serve as intermediate hosts thrive in ponds, rice paddies, and the marshy areas that develop when dams are built. Tapeworms inhabit the intestine of vertebrates Adult members of the more than 5000 species of class Cestoda live as parasites in the intestine of probably every kind of vertebrate, including humans. Tapeworms are long, flat, ribbonlike animals strikingly specialized for their parasitic mode of life. Among their many adaptations are suckers and sometimes hooks on the “head,” or scolex,

2

Larvae burrow through skin.

Courtesy CDC

1

Larvae make their way to circulatory system, where they mature. During reproduction, which takes place in veins, male holds female in a long groove (as shown).

1 mm

7

3

Finally, fork-tailed larvae (cercariae) develop and leave snail.

4

6

Larvae must enter a second host, a freshwater snail. After burrowing into tissues of snail, larvae develop into a form that reproduces asexually. This process greatly increases the number of larvae.

FIGURE 31-9 Animated Life cycle of the blood fluke (Schistosoma) The LM shows an adult male enfolding a smaller female.

5

Eggs pass into intestine.

Eggs containing developing embryos are excreted with feces.

If they find their way to fresh water, the eggs hatch, releasing free-swimming larvae (miracidia).

tapeworms have complex life cycles, spending their larval stage within the body of an intermediate host and their adult life within the body of a different, final host. FIGURE 31-11 illustrates the life cycle of the beef tapeworm, which can infect humans when they eat undercooked beef containing the larvae.

© SPL/Photo Researchers, Inc.

Nemerteans are characterized by their proboscis

250 μm

FIGURE 31-10 Scolex of a tapeworm The small tapeworm (Acanthrocirrus retrisrostris) reaches maturity in the intestine of wading birds that eat barnacles. The color-enhanced SEM shows the pistonlike cluster of hooks that can be withdrawn into the head or thrust out and buried in the host’s tissue. You can see two of the four powerful suckers beneath the hooks.

The nemerteans (neh-mur¿-tee-uns; phylum Nemertea) include the ribbon worms, a relatively small group of about 1200 species of free-living animals (FIG. 31-12). Most burrow in marine sediments, but a few species inhabit deep sea water, fresh water, or damp soil. Many nemerteans are carnivorous, feeding on crustaceans and annelids. Nemerteans have long, narrow bodies, either cylindrical or flattened, ranging in length from 1 mm to more than 30 m (100 ft). Some are a vivid orange, red, or green, with black or colored stripes. Their most remarkable organ, the proboscis, is a long, hollow, muscular tube that can be rapidly everted (turned inside out) from the anterior end of the body. Used to capture prey, the sticky proboscis can be wrapped around a small animal. In some species the proboscis is sharp, and in various species it secretes toxic fluid that immobilizes prey. The proboscis is a derived character that distinguishes nemerteans from all other invertebrate groups. Because of this character, these animals are sometimes called proboscis worms. Historically, the nemerteans were of special evolutionary interest because biologists thought they were the earliest animals to have a circulatory system and a complete digestive tract. Nemerteans have no heart; blood is circulated by contractions of muscular blood vessels and by movements of the body. 2

that enable the parasite to attach to the host’s intestine (FIG. 31-10). The reproductive adaptations and abilities of tapeworms are extraordinary. The body of the tapeworm consists of a long chain of segments called proglottids. Each proglottid is an entire reproductive machine equipped with both male and female reproductive organs. A single proglottid contains up to 100,000 eggs. Because an adult tapeworm may have as many as 2000 segments, its reproductive potential is staggering. A single tapeworm can produce 600 million eggs per year! Proglottids farthest from the tapeworm’s head contain the ripest eggs; these segments are shed from the host’s body along with the feces. The tapeworm has no mouth or digestive system. Digested food from the host is absorbed across the worm’s body wall. The tapeworm also lacks well-developed sense organs. Some

1

Magnified view of muscle containing encysted larvae. 3

Cattle ingest zygotes with grass. Larvae hatch and encyst in muscle of cow or steer.

When human eats poorly cooked, infected beef, digestive juices dissolve the cyst, freeing larva.

Encysted larvae

4

Zygotes

6

Magnified view of zygotes on grass. 5

Mature proglottids leave body with feces and burst open, releasing zygotes.

FIGURE 31-11 Animated Life cycle of the beef tapeworm, Taenia saginata

After attaching itself to lining of human small intestine, larva matures into adult tapeworm. Zygotes are produced within proglottids.

Deuterostomia Kjell B. Sandved

Choanoflagellate ancestor

Ecdysozoans

Rotifers

Lophophorates

Annelids

Mollusks

Nemerteans

Platyhelminthes

Cniderians

Ctenophores

Sponges

Lophotrochozoa

(a) Ribbon worm (Lineus) from the Pacific coast of Panama.

Cerebral ganglion

Rhynchocoel Retractor muscle

Anus Mouth Extended proboscis

Digestive tract

(b) Lateral view of a typical nemertean. Note the complete digestive tract that extends from mouth to anus, giving this animal a tube-within-a-tube body plan.

FIGURE 31-12 Nemerteans (ribbon worms)

The chamber surrounding the proboscis is a coelomic space, known as a rhynchocoel. It develops in the embryo as the coelom does in coelomate protostomes (see Chapter 30). However, it differs in position and in function from the coelom of other animals. Biologists now view the rhynchocoel as a remnant of the coelom of a coelomate ancestor. Because of its presence and recent molecular evidence, biologists now classify nemerteans with the lophotrochozoans. Like the flatworms, nemerteans evolved from coelomate animals and became simpler over time.

Mollusks have a muscular foot, visceral mass, and mantle The mollusks (phylum Mollusca) include clams, oysters, snails, slugs, octopods, and the largest of all the invertebrates, the giant squid, which averages 9 to 16 m (about 30 to 53 ft) in length, including its tentacles. More than 80,000 living species and 35,000 fossil species (second only to the arthropods in number) have been described. Here we discuss four of eight recognized classes. We illustrate representatives of these classes in FIGURE 31-13 and describe them in TABLE 31-3. Although most mollusks are marine, many snails and clams live in fresh water, and some species of snails and slugs inhabit the land. Mollusks probably evolved early in the history of the proto-

stome clade—soon after the evolution of the coelom, but before the origin of the segmented body that is characteristic of annelids. Although mollusks vary widely in outward appearance, most share six basic characteristics: 1. A soft body, usually covered by a dorsal shell composed mainly of calcium carbonate. 2. A broad, flat, muscular foot, located ventrally, which is used for locomotion. 3. The body organs (viscera) concentrated as a visceral mass located above the foot. 4. The dorsal body wall forms a pair of folds called the mantle. The mantle is a thin sheet of tissue that generally overhangs the visceral mass, forming a mantle cavity. The mantle may contain glands that secrete a shell. The mantle cavity contains gills or a lung. 5. A rasplike structure called a radula, which is a belt of teeth in the mouth region. (The radula is not present in clams or their relatives, which are filter feeders.) 6. A coelom, generally reduced to small compartments around certain organs, including the heart and excretory organs (metanephridia). The main body cavity is typically a hemocoel, a space containing blood (see following discussion of open circulatory systems). The hemocoel is not a coelom.

TABLE 31-3

Major Classes of Phylum Mollusca

Class and Representative Animals

Characteristics

Polyplacophora Chitons Gastropoda Snails, slugs, nudibranchs Bivalvia Clams, oysters, mussels Cephalopoda Squids, octopods

Marine; dorsal shell consisting of eight separate transverse plates; head reduced; broad foot used for locomotion Marine, freshwater, or terrestrial; coiled shell in many species; torsion of visceral mass; well-developed head with tentacles and eyes Marine or freshwater; body laterally compressed; two-part shell hinged dorsally; hatchet-shaped foot; filter feeders Marine; predatory; foot modified into tentacles, usually bearing suckers; well-developed eyes; closed circulatory system

Kjell B. Sandved

Deuterostomia

Ecdysozoans

Rotifers

Lophophorates

Annelids

Mollusks

Nemerteans

Platyhelminthes

Cniderians

Ctenophores

Sponges

Lophotrochozoa

Shell Digestive tract

(a) Class Polyplacophora

Shell

(b) Class Gastropoda

The broad, flat foot of the gastropod is an adaptation to its mobile lifestyle. The mystery snail (Pomacea bridgesi) inhabits fresh water and can be found burrowing in the mud.

Internal shell

Tentacles (modified foot)

Digestive tract

(d) Class Cephalopoda

FIGURE 31-13 Animated The mollusk body plan

Digestive Foot tract

(c) Class Bivalvia

The compressed body of the horseneck clam (Tresus capax) is adapted for burrowing in the North Pacific mud. Its shell grows to about 20 cm (8 in) in length.

Jeffrey Torman/Peter Arnold, Inc.

Foot

Digestive tract

Shell Tom McHugh/Photo Researchers, Inc.

E. R. Degginger/Dembinsky Photo Associates

Choanoflagellate ancestor

Chitons are sluggish marine animals with shells composed of eight overlapping plates. This sea cradle chiton (Tonicella lineata), which reaches about 5 cm (2 in) in length, inhabits rocks covered with coralline algae in coastal waters off the U.S. Pacific Northwest.

The squid body is streamlined for swimming. To avoid being seen by potential predators, the squid can change color to blend with its background. This northern short-fin squid (Illex illecebrosus), which grows to a mantle length of up to about 31 cm (about 1 ft), inhabits the North Atlantic Ocean.

Mollusks have all of the organ systems typical of complex animals. The digestive system is a tube, often coiled, consisting of a mouth, buccal cavity (mouth cavity), esophagus, stomach, intestine, and anus. The mollusk radula, located within the buccal cavity, projects out of the mouth and is used to scrape particles of food from the surface of rocks or the ocean floor. Sometimes the radula is used to drill a hole in the shell of a prey animal or to tear off pieces of a plant. Most mollusks have an open circulatory system in which the blood, called hemolymph, bathes the tissues directly. The heart pumps blood into a single blood vessel, the aorta, which may branch into other vessels. Eventually, blood flows into a network of large spaces called sinuses, bringing the blood into direct contact with the tissues. This network makes up the hemocoel, or blood cavity. From the sinuses, blood drains into vessels that conduct it to the gills, where it is recharged with oxygen. After passing through the gills, the blood returns to the heart. Thus, blood flow in a mollusk follows this pattern: heart ¡ aorta ¡ smaller blood vessels ¡ blood sinuses (hemocoel) ¡ blood vessels to gills ¡ heart

In open circulatory systems, blood pressure tends to be low, and tissues are not very efficiently oxygenated. Because most mollusks are slow-moving animals with low metabolic rates, this type of circulatory system is adequate. The active cephalopods (the class that includes the squids and octopods) have a closed circulatory system in which blood flows through a complete circuit of blood vessels. Most marine mollusks pass through one or more larval stages. The first stage is typically a trochophore larva, a free-swimming, top-shaped larva with two bands of cilia around its middle (FIG. 31-14). In many mollusks (gastropods such as snails and

Digestive tract

bivalves such as clams), the trochophore larva develops into a veliger larva, which has a shell, foot, and mantle. The veliger larva is unique to the mollusks. Chitons may be similar to ancestral mollusks Polyplacophorans (pol-ee-plah-kof¿-o-rans; meaning “many plates”) are chitons, marine animals with flattened bodies (see Fig. 31-13a). Their most distinctive feature is a shell composed of eight separate but overlapping dorsal plates. The head is reduced, and there are no eyes or tentacles. The chiton inhabits rocky intertidal zones, using its broad, flat foot to move and to hold on firmly to rocks. By pressing its mantle against the substratum and lifting the inner edge of the mantle, the chiton can produce a partial vacuum. The resulting suction lets the animal adhere powerfully to its perch. Using its radula for grazing, the chiton scrapes algae and other small organisms off rocks and shells. Gastropods are the largest group of mollusks The gastropods, the snails, slugs, conchs, sea slugs, and their relatives, are the largest and most diverse group of mollusks (see Fig. 31-13b). In fact, with more than 70,000 extant species, gastropods constitute the second-largest class in the animal kingdom, second only to insects. Most gastropods inhabit marine waters, but others make their homes in brackish or fresh water or on land. We think of snails as having a single, spirally coiled shell into which they can withdraw the body, and many do. However, other gastropods, such as limpets, have shells like flattened dunce caps. Still others, such as garden slugs and the beautiful marine slugs known as nudibranchs, have no shell at all (FIG. 31-15). Many gastropods have a well-developed head with tentacles. Two simple eyes may be located on stalks that extend from the head. The gastropod uses its broad, flat foot for creeping. Most land snails do not have gills. Instead, the mantle is highly vascularized and functions as a lung. Biologists describe these garden snails and slugs as pulmonate (“having a lung”).

Cilia

Mouth © Bruce Watkins/Animals Animals

Nephridium Mesodermal cells Anus

FIGURE 31-14 A trochophore larva The first larval stage of a marine mollusk, the trochophore larva, is also characteristic of annelids. Just above the mouth a band of ciliated cells functions as a swimming organ and, in some species, collects suspended food particles.

FIGURE 31-15 Nudibranch A nudibranch (Chromodoris porterae) feeds on a lightbulb tunicate (Clavelina huntsmani).

Shell Anus

Mantle cavity

Mouth

FIGURE 31-16 Animated Torsion in a gastropod As the bilateral larva develops, the visceral mass twists 180 degrees relative to the head. The digestive tube coils, and the anus becomes relocated near the mouth.

Torsion, a twisting of the visceral mass, is a unique feature of gastropods. This twisting is unrelated to the coiling of the shell. As the bilateral larva develops, one side of the visceral mass grows more rapidly than the other side. This uneven growth rotates the visceral mass. The visceral mass and mantle twist permanently up to 180 degrees relative to the head. As a result, the digestive tract becomes somewhat U shaped, and the anus comes to lie above the head and gill (FIG. 31-16). Subsequent growth is dorsal and usually in a spiral coil. Torsion limits space in the body, and typically the gill, excretory organ (metanephridium), and gonad are absent on one side. Some biologists hypothesize that torsion is an adaptation that protects the head by allowing it to enter the shell first during withdrawal from potential predators. Without torsion, the foot would be withdrawn first.

slowly through rock or wood, seeking protected dwellings. The shipworm, Teredo, a clam that damages dock pilings and other marine installations, is just looking for a home. A few bivalves, such as scallops, swim rapidly by clapping their two shells together with the contraction of a large adductor muscle (the part of the scallop that humans eat). Clams and oysters are filter feeders that trap food particles suspended in sea water. They take water in through an extension of the mantle called the incurrent siphon. Water leaves by way of an excurrent siphon. As the water passes over the gills, mucus secreted by the gills traps food particles in the stream of water. Cilia move the food to the mouth. An oyster can filter more than 30 L (about 32 qt) of water per hour! As filter feeders, bivalves have no need for a radula, and indeed they are the only group of mollusks that lack this structure. Cephalopods are active predators In contrast with most other mollusks, members of class Cephalopoda (meaning “head-foot”) are fast-swimming predators (see Fig. 31-13d). The cephalopod mouth is surrounded by tentacles, or arms: 8 in octopods, 10 in squids, and as many as 90 in the chambered nautilus. The large head has well-developed eyes that form images. Although they develop differently, their complex eyes are structurally similar to vertebrate eyes and function in much the same way. The octopus has no shell; and the shell of the squid, located inside the body, is greatly reduced. Nautilus has a coiled shell consisting of many chambers built up over time. Each year the animal lives in the newest and largest chamber of the series. Nautilus secretes a mixture of gases similar to air into the other chambers. By regulating the amount of gas in the chambers, the animal controls its depth in the ocean. The tentacles of squids, octopods, and cuttlefish are covered with suckers for seizing and holding prey. In addition to a radula,

Most bivalves are filter feeders The bivalves include clams, oysters, mussels, scallops, and their relatives (see Fig. 31-13c). Typically, the soft body is laterally compressed and completely enclosed by a two-part shell that hinges dorDigestive gland sally and opens ventrally (FIG. 31-17). This arDORSAL rangement allows the hatchet-shaped foot Stomach to protrude ventrally for locomotion and for burrowing in mud. The two parts, or valves, of Esophagus the shell are connected by an elastic ligament. Ganglion Stretching of the ligament opens the shell. Large, strong adductor muscles attached to the Anterior adductor shell permit the animal to close its shell. muscle The inner, pearly layer of the bivalve shell Mouth is made of calcium carbonate secreted in thin sheets by the epithelial cells of the mantle. Palp Known as mother-of-pearl, this material is valPedal ued for making jewelry and buttons. Should a ganglion bit of foreign matter lodge between the shell and the mantle, the epithelial cells covering Foot Intestine Gonad the mantle secrete concentric layers of calcium carbonate around the intruding particle. VENTRAL Many species of oysters and clams form pearls in this way. FIGURE 31-17 Animated Internal anatomy of a clam Some bivalves, such as oysters, attach The two shells of a bivalve hinge dorsally and open ventrally. permanently to the substrate. Others burrow

Heart Metanephridium Intestine Posterior adductor muscle Anus Excurrent siphon Incurrent siphon Gill—partially cut Mantle Shell

the mouth has two strong, horny beaks used to kill prey and tear it to bits. The thick, muscular mantle is fitted with a funnel-like siphon. By filling the cavity with water and ejecting it through the siphon, the cephalopod achieves forceful jet propulsion. Squids and cuttlefish have streamlined bodies adapted for efficient swimming. In addition to speed, the cephalopod has two other adaptations that facilitate escape from its predators, which include certain whales and moray eels. It can confuse the enemy by rapidly changing colors. By expanding and contracting chromatophores, cells in the skin that contain pigment granules, the cephalopod can display an impressive variety of mottled colors. Another defense mechanism is its ink sac, which produces a thick, black liquid the animal releases in a dark cloud when alarmed. While its enemy pauses, temporarily blinded and confused, the cephalopod escapes. The ink inactivates the chemical receptors of some predators, making them incapable of detecting their prey. The octopus feeds on crabs and other arthropods, catching and killing them with a poisonous secretion of its salivary glands. During the day the octopus usually hides among the rocks; in the evening it emerges to hunt for food. Its motion is incredibly fluid, giving little hint of the considerable strength in its eight arms. Small octopods survive well in aquariums and have been studied extensively. Because octopods are relatively intelligent and can make associations among stimuli, researchers have used them as models for studying learning and memory. Their highly adaptable behavior more closely resembles that of the vertebrates than the more stereotypic patterns of behavior observed in other invertebrates (see Chapter 52).

Annelids are segmented worms Annelids (an¿-eh-lids; phylum Annelida) are segmented worms with bilateral symmetry and a tubular body that may be partitioned into more than 100 ringlike segments. This phylum, composed of

about 15,000 species, includes three main groups: the polychaetes, a group of marine and freshwater worms; the earthworms; and the leeches (TABLE 31-4 and FIG. 31-18). The term Annelida (from a Latin word meaning “little rings”) refers to the series of rings, or segments, which make up the annelid body. Both the body wall and many of the internal organs are segmented. Some structures, such as the digestive tract and certain nerves, extend the length of the body, passing through successive segments. Other structures, such as excretory organs, are repeated in each segment. In polychaetes and earthworms, segments are separated from one another internally by transverse partitions called septa. An important advantage of segmentation is that it facilitates locomotion. The coelom is divided into segments, and each segment has its own muscles. This arrangement allows the animal to elongate one part of its body while shortening another part. The annelid’s hydrostatic skeleton is important in movement. Polychaetes and earthworms have bristlelike structures called setae (sing., seta), located on each segment. Setae provide traction as the worm moves TABLE 31-4 Class and Representative Animals Polychaeta Sandworms, tubeworms Oligochaeta Earthworms Hirudinida Leeches

Major Classes of Phylum Annelida

Characteristics Mainly marine; each segment bears a pair of parapodia with many setae; well-developed head; separate sexes; trochophore larva Terrestrial and freshwater worms; few setae per segment; lack well-developed head; hermaphroditic Most are blood-sucking parasites that inhabit fresh water; appendages and setae absent; prominent muscular suckers

Deuterostomia

Ecdysozoans

Rotifers

Lophophorates

Annelids

Mollusks

Nemerteans

Platyhelminthes

Cniderians

Ctenophores

Sponges

Lophotrochozoa

Setae

Mouth

Clitellum

Suckers

Parapodia Choanoflagellate ancestor

Anus

(a) Class Polychaeta. Polychaetes are marine worms with paddleshaped parapodia.

FIGURE 31-18 Three major groups of annelids

(b) Class Oligochaeta. Oligochaetes, which include the earthworms, inhabit freshwater and moist terrestrial areas.

(c) Class Hirudinida. Many leeches are parasites that feed on blood.

along by alternating contraction of its longitudinal and circular muscles. In earthworms, setae consist mainly of chitin. The annelid nervous system typically consists of a ventral nerve cord and a simple brain consisting of a pair of ganglia. Each segment has a pair of ganglia and lateral nerves. Annelids have a large, well-developed coelom, a closed circulatory system, and a complete digestive tract extending from mouth to anus. Respiration is cutaneous, that is, through the skin, or by gills. Typically, a pair of excretory tubules called metanephridia is located in each segment (described in Chapter 48).

also be equipped with tentacles and palps (feelers). Polychaetes develop from free-swimming trochophore larvae similar to those of mollusks. Behavioral patterns that ensure fertilization have evolved in many polychaete species. By responding to certain rhythmic variations, or cycles, in the environment, nearly all the females and males of a given species release their gametes into the water at the same time. For example, more than 90% of reef-dwelling Palolo worms of the South Pacific shed their eggs and sperm within a 2-hour period on one night of the year. The posterior portion of the Palolo worm, loaded with eggs or sperm, breaks off from the rest of the body. It comes to the surface and bursts, releasing its gametes. This mass reproductive event occurs in October or November at the beginning of the last lunar quarter. Local islanders gather great numbers of the swarming polychaetes, which they bake or eat raw.

Most polychaetes are marine worms with parapodia Most polychaetes are marine worms, and many swim freely in the sea. Some polychaetes are part of the plankton and are important in marine food chains. Others burrow in the mud near the shore or in crevices of rocks or coral reefs. Still others live in tubes they secrete or make by cementing bits of shell and sand together with mucus. Giant tube worms have been found in deep vents in the floor of the Pacific Ocean (see Inquiring About: Life Without the Sun in Chapter 55). Each body segment typically bears a pair of paddle-shaped appendages called parapodia (sing., parapodium) that function in locomotion and in gas exchange. These fleshy structures bear many stiff setae (the term polychaete means “many bristles”; see the fireworm in the chapter opening photograph). Most polychaetes have a well-developed head with eyes and antennae. The head may

Earthworms help maintain fertile soil The approximately 3100 species of oligochaetes live almost exclusively in fresh water and in moist terrestrial habitats. These worms lack parapodia, have few bristles per segment (the term oligochaete means “few bristles”), and lack a well-developed head. All oligochaetes are hermaphroditic. Lumbricus terrestris, the common earthworm, is about 20 cm (8 in) long. Its body has more than 100 segments, separated externally by grooves that indicate the internal position of the septa (FIG. 31-19). The earthworm’s body is somewhat protected from

Clitellum

Intestine Intestine Gizzard

Crop

Dorsal blood vessel Metanephridia

Esophagus Pharynx

Dorsal vessel Ventral vessel Muscles

Longitudinal Circular

Nerve cord

Cerebral ganglia

Septa

(a) The internal structure has been exposed at the anterior end of an earthworm.

FIGURE 31-19 Animated Body plan of an earthworm

Mouth Coelom Ventral blood vessel

Nerve cord

(b) Cross section of an earthworm.

drying by a thin, transparent cuticle secreted by the cells of the epidermis. Mucus secreted by gland cells of the epidermis forms an additional protective layer over the body surface. The body wall has an outer layer of circular muscles and an inner layer of longitudinal muscles. An earthworm literally eats its way through the soil. It ingests its own weight in soil and nutritious decaying vegetation every 24 hours! Earthworms enhance the formation and maintenance of fertile soil. As they move about, they turn and aerate the soil and enrich it with nitrogenous wastes. The earthworm processes its meal of soil in a complex digestive system. Food is swallowed through the muscular pharynx and passes through the esophagus, which is modified to form a thin-walled crop where food is stored, and a thick-walled, muscular gizzard. In the gizzard, food is ground to bits by sand grains consumed along with the meal. The rest of the digestive system is a long, straight intestine that chemically digests food and absorbs nutrients. Wastes pass out of the intestine to the exterior through the anus. The efficient closed circulatory system consists of two main blood vessels that extend longitudinally. The dorsal blood vessel, just above the digestive tract, collects blood from vessels in the segments, then contracts to pump the blood anteriorly. In the region of the esophagus, five pairs of blood vessels propel blood from the dorsal to the ventral blood vessel. Located just below the digestive tract, the ventral blood vessel carries blood posteriorly. Small vessels branch from it and deliver blood to the various structures in each segment, as well as to the body wall. Within these structures, blood flows through tiny capillaries before returning to the dorsal blood vessel. Gas exchange takes place through the moist skin. Oxygen is transported by hemoglobin, a respiratory pigment in the blood. The excretory system consists of paired metanephridia, repeated in almost every segment of the body. The nervous system consists of a pair of cerebral ganglia (a simple brain) just above the pharynx and a subpharyngeal ganglion just below the pharynx. A ring of nerve fibers connects these ganglia. From the lower ganglion, a ventral nerve cord extends beneath the digestive tract to the posterior end of the body. A pair of fused ganglia is present in each segment along the nerve cord. Nerves extend laterally from the ganglia to the muscles and other structures of that segment. The ganglia coordinate muscle contractions of the body wall, allowing the worm to creep along. Like other oligochaetes, earthworms are hermaphroditic. During copulation, two worms, headed in opposite directions, press their ventral surfaces together. These surfaces become glued together by the thick mucous secretions of each worm’s clitellum, a thickened ring of epidermis. Sperm are then exchanged and stored in the seminal receptacles (small sacs) of the other worm. A few days later, each clitellum secretes a membranous cocoon containing a sticky fluid. As the cocoon is slipped forward, eggs are laid in it. Sperm are added as the cocoon passes over the openings of the seminal receptacles. As the cocoon slips free over the worm’s head, its openings constrict so that a spindle-shaped capsule is formed. The fertilized eggs develop into tiny worms within this capsule. This reproductive pattern is an adaptation to terrestrial

life; the cocoon protects the delicate gametes and young worms from drying out. Many leeches are blood-sucking parasites Leeches are members of a group called Hirudinida. Most leeches inhabit fresh water, but some live in the sea or in moist areas on land. Some leeches are nonparasitic predators that capture small invertebrates such as earthworms and snails. However, about 75% of the known species of leeches are blood-sucking parasites. Leeches differ from other annelids in having neither setae nor parapodia. Leeches have muscular suckers at both the anterior and posterior ends of their body. Most parasitic leeches attach themselves to a vertebrate host, bite through the skin, and suck out a quantity of blood, which is stored in pouches in the digestive tract. Hirudin, an anticoagulant secreted by glands in the crop, ensures leeches a full meal of blood. In 31 minutes, a leech can suck out as much as 10 times its own weight in blood! Some leeches feed only about twice each year because they digest their food slowly over several months. Leeches have been used since ancient times for drawing blood from areas swollen by poisonous stings and bites. During the 19th century, they were widely used to remove “bad blood,” thought to be the cause of many diseases. Leeches are used in modern medicine to remove excess fluid and blood that accumulate within body tissues as a result of injury, disease, or surgery (FIG. 31-20). The leech attaches its sucker near the site of injury, makes an incision, and secretes hirudin. The hirudin prevents the blood from clotting and dissolves clots that already exist.

The lophophorates are distinguished by a ciliated ring of tentacles With a few exceptions, lophophorates are marine animals adapted for life on the ocean floor. These coelomates are distinguished by their lophophore, a ring of ciliated tentacles that surrounds the mouth. The lophophore is an adaptation for capturing suspended particles in the water. Lophophorates do not have a distinct head. The three lophophorate groups are Brachiopoda, Phoronida, and Bryozoa (also known as Ectoprocta). Brachiopods, or lampshells, are solitary marine animals that inhabit cold water. Until the mid-19th century, they were thought to be mollusks because they are suspension feeders that superficially resemble clams and other bivalve mollusks. Their body is enclosed between two shells and has a mantle and mantle cavity (FIG. 31-21a). However, they differ from bivalve mollusks in that the shells are dorsal and ventral rather than lateral; each shell is symmetrical about the midline, and the two shell valves are typically of unequal size. Brachiopods attach to the substrate by a long stalk. The action of cilia on the lophophore brings water with suspended food into the slightly opened shell. Although there are now only about 325 living species, brachiopods were abundant and diversified during the Paleozoic era. Paleobiologists have described about 12,000 fossil species. Only about 20 extant species of phoronids are known. They are wormlike, sessile animals found in coastal marine sediments. Adult phoronids secrete chitinous tubes in which they live (FIG. 31-21b).

St. Bartholomew’s Hospital/Science Photo Library/Photo Researchers, Inc.

T. E. Adams/Visuals Unlimited

500 μm

(a) LM of a blood-sucking leech (Helobdella stagnalis) that feeds on mammals. The digestive tract (dark area) of its swollen body is filled with ingested blood.

(b) Medicinal leeches (Hirudo medicinalis) are used here to treat hematoma, an accumulation of blood within tissues that results from injury or disease.

FIGURE 31-20 Leeches

Andrew J. Martinez / Photo Researchers, Inc.

Deuterostomia

Ecdysozoans

Rotifers

Lophophorates

Annelids

Mollusks

Nemerteans

Platyhelminthes

Ctenophores

Cniderians

Sponges

Lophotrochozoa

(a) Brachiopods. Like other brachiopods, these northern lampshells (Terebratulina septentrionalis) superficially resemble clams.

M. I. Walker /Photo Researchers, Inc.

Choanoflagellate ancestor

Stan Elems /Visuals Unlimited

They extend their lophophores from their tubes for feeding. Phoronids vary in length from 2 cm to more than 20 cm (less than 1 in to about 8 in), but they are only 1 to 3 mm in diameter. Bryozoans, also known as ectoprocts or “moss animals,” are microscopic aquatic animals. They form sessile colonies by asexual budding (FIG. 31-21c). Each colony can consist of millions of individuals and can extend in length up to about 31 cm (about 1 ft). The colonies typically appear plantlike, but some have the appearance of coral. Approximately 4500 living species are known. Biologists are still debating the evolutionary position of the lophophorates. They were traditionally considered a monophyletic group and a sister group to the deuterostomes, based on certain structural similarities. However, these lophophorate groups also have several morphological characters that suggest a relationship with the lophotrochozoans. Although their phylogeny is still

(b) Phoronids. A single lophophore of a phoronid (Phoronopsis viridis). This animal is about 20 cm (8 in) long, but only 1–3 mm in diameter.

FIGURE 31-21 Representative lophophorates

(c) Bryozoans. Like many bryozoans, this freshwater species (Cristatella mucedo) forms colonies by asexual budding.

being debated, based on molecular data (18S rDNA nucleotide sequences), the lophophorate phyla are currently classified as Lophotrochozoa.

Rotifers have a crown of cilia Rotifers (phylum Rotifera), or “wheel animals,” are among the less familiar invertebrates. Although no larger than many protozoa, these aquatic, microscopic animals are multicellular. Most of the 2000 described species inhabit fresh water, but some live in marine environments or damp soil. Rotifers are important in cycling nutrients in aquatic ecosystems. Rotifers do not have a true coelom and were formerly classified as pseudocoelomates. Current evidence suggests that rotifers evolved from animals with a true coelom and we classify them as lophotrochozoans. Rotifers have a characteristic crown of cilia on their anterior end. The cilia beat rapidly during swimming and feeding, giving the appearance of a spinning wheel (FIG. 31-22). Rotifers feed on tiny organisms suspended in the stream of water drawn into the mouth by the cilia. A muscular organ posterior to the mouth grinds the food. The complete digestive tract ends in an anus through which wastes are eliminated. Rotifers have a nervous system with a “brain” and sense organs, including eyespots. Protonephridia with flame cells remove excess water from the body and may also excrete metabolic wastes.

Biologists have discovered that rotifers, like some other animals with a pseudocoelom, are “cell constant.” Each member of a given species has exactly the same number of cells. Indeed, each part of the body has a precisely fixed number of cells arranged in a characteristic pattern. Cell division does not take place after embryonic development, and mitosis cannot be induced. Growth results from an increase in cell size. Do you think rotifers could develop cancer? One of the challenging research problems of our time is to discover the difference between such nondividing cells and the dividing cells of other animals. Another interesting characteristic of rotifers is their ability to survive very dry conditions and temperature extremes. These interesting animals can remain in a dormant state for months or years. When conditions become favorable, they recover and become active once again.

Review ■

■ ■

■

What are some advantages of (a) bilateral symmetry, (b) cephalization, and (c) a coelom? On what basis have biologists classified nemerteans as lophotrochozoans? How does the lifestyle of a gastropod differ from that of a cephalopod? Identify adaptations that have evolved in each for its particular lifestyle. What are two distinguishing characteristics for each of the following: (1) flatworms, (2) mollusks, and (3) annelids?

Crown of cilla Deuterostomia

Ecdysozoans

Rotifers

Lophophorates

Annelids

Mollusks

Nemerteans

Platyhelminthes

Cniderians

Ctenophores

Sponges

Lophotrochozoa

Mouth Eyespot

Pharynx (grinding organ) Digestive gland

John Walsh /Science Photo Library / Photo Researchers, Inc.

Choanoflagellate ancestor

Stomach Reproductive organ

Intestine

Protonephridium

Bladder Anus

Cuticle and epidermis

150 μm

(a) LM of an Antarctic rotifer (Philodina gregaria) that survives the winter by forming a cyst. It reproduces in great numbers, sometimes coloring the water red.

FIGURE 31-22 Rotifers (wheel animals)

(b) Longitudinal section showing rotifer anatomy. The motion of its cilia draws particles of food such as algae into the mouth.

tems; however, they lack specific circulatory structures. The sexes are usually separate, and the male is generally smaller than the female. Traditionally, biologists classified nematodes as a single phylum based on the pseudocoelom, wormlike body structure, and other characters such as absence of a well-defined head. However, molecular evidence suggests that nematodes are not a monophyletic group and that they evolved from more complex animals by a simplification in body plan. Like arthropods and other ecdysozoans, the nematode sheds its cuticle periodically. Systematists currently classify nematodes with the Ecdysozoa (see Fig. 30-7). Although most nematodes are free-living, others are important parasites in plants and animals. More than 50 species of roundworms are human parasites, including Ascaris, hookworms, pinworms, and the trichina worm. The common intestinal parasite Ascaris spends its adult life in the human intestine, where it ingests partly digested food (FIG. 31-24). Ascaris is a white worm about 25 cm (10 in) long. Like most parasites, it has remarkable reproductive adaptations that ensure survival of its species. A mature female may produce as many as 200,000 eggs per day! Hookworms attach to the lining of the intestine and suck blood, potentially causing serious tissue damage and blood loss. Pinworms are the most common worms found in children. The tiny pinworm eggs are often ingested by eating with hands contaminated with them. The trichina worm lives inside a variety of animals, including pigs, rats, and bears. Humans typically become infected by eating undercooked, infected meat. Larvae encyst in skeletal muscle.

31.3 THE ECDYSOZOA ■ ■ LEARNING OBJECTIVES 9 Summarize what is known about the phylogeny of the ecdysozoans. 10 Describe distinguishing characteristics of nematodes, and identify four parasitic nematodes.

11 Describe six key characteristics of arthropods, summarize proposed phylogeny of arthropods, and distinguish among arthropod subphyla and classes; give examples of animals that belong to each group. 12 Identify adaptations that have contributed to the biological success of insects.

The Ecdysozoa include the nematodes (roundworms) and arthropods, animals characterized by a cuticle, a noncellular body covering secreted by the epidermis. The name Ecdysozoa refers to the process of ecdysis, or molting, characteristic of animals in this group. During ecdysis, an animal sheds its outer covering (for example, cuticle, exoskeleton, skin), which is then replaced by the growth of a new one. The taxonomic validity of the Ecdysozoa has been supported by many types of data, including molecular data.

Roundworms are of great ecological importance

T. E. Adams / Visuals Unlimited

Deuterostomia

Arthropods

Nematodes

Lophotrochozoans

Ctenophores

Cnidarians

Sponges

Nematodes (nem¿-uh-todes; phylum Nematoda), or roundworms, play key ecological roles as decomposers and predators of smaller organisms. Many soil nematodes eat bacteria. Nematodes are numerous and widely distributed in soil and in marine and freshwater sediments. About 80,000 species have been described, and many more remain to be discovered. The nematodes rival the insects in number of individuals. A spadeful of soil may contain thousands of these mainly microscopic worms, which thrash Arthropods are characterized by jointed around, coiling and uncoiling. Greater understanding of soil nemappendages and an exoskeleton of chitin atode diversity and ecology will contribute to solving some imporArthropods (ar¿-thro-pods; phylum Arthropoda) are the most tant agricultural problems. Caenorhabditis elegans, a free-living soil biologically successful group of animals. More than 80% of all nematode, is an important model research organism for biologists known animals are arthropods! They are more diverse and live in a studying the genetic control of development (see Chapter 17). The elongated, cylindrical, threadlike nematode body is pointed Ecdysozoa at both ends and covered with a tough, flexible cuticle (FIG. 31-23). Secreted by the underlying epidermis, the thick cuticle gives the nematode body shape and offers some protection. The epidermis is unusual in that it does not consist of distinct cells. Nematodes have a fluid-filled pseudocoelom that serves as a hyChoanoflagellate drostatic skeleton. It transmits the ancestor force of muscle contraction to the enclosed fluid. Movement of fluid in the pseudocoelom is also important in nutrient transport and distribu250 μm tion. Like ribbon worms, nematodes exhibit bilateral symmetry, a comFIGURE 31-23 LM of a free-living nematode plete digestive tract, three definite This unidentified aquatic nematode is shown among the cyanobacterium Oscillatoria, which it eats. tissue layers, and definite organ sys-

Mouth

Pharynx

Dorsal nerve Excretory canal

Pharynx Muscle of pharynx wall

Excretory gland

Pseudocoelom Uterus Ovary Intestine Excretory canal

Vulva Muscle layer

Ventral nerve

Epidermis Cuticle

(b) Cross section

Anus

(a) Longitudinal section

FIGURE 31-24 Animated The roundworm Ascaris (a) Note the complete digestive tract that extends from mouth to anus. (b) This cross section through Ascaris shows the tube-within-a-tube body plan. The protective cuticle that covers the body helps the animal resist drying.

greater range of habitats than do the members of any other animal phylum. More than one million species have been described, and biologists predict millions more will be identified. The following adaptations have greatly contributed to their success: 1. The arthropod body, like that of the annelid, is segmented. Segmentation is important from an evolutionary perspective because it provides the opportunity for specialization of body regions. In arthropods, groups of segments are specialized to perform particular functions. Segments differ in shape, muscles, or the appendages they bear. 2. A hard exoskeleton, composed of chitin and protein, covers the entire body and appendages. The exoskeleton serves as a coat of armor that protects against predators and helps prevent excessive loss of moisture. It also supports the underlying soft tissues. Arthropods move effectively because distinct muscles attach to the inner surface of the exoskeleton and operate the joints of the body and appendages.

A disadvantage of the exoskeleton is that it is nonliving, and the arthropod periodically outgrows it. Recall that molting is the process of shedding an old exoskeleton, which is then replaced by a larger one. The shed exoskeleton represents a net metabolic loss, and molting also leaves the arthropod temporarily vulnerable to predators. 3. Paired, jointed appendages, from which this group gets its name (arthropod means “jointed foot”), are modified for many functions. They serve as swimming paddles, walking legs, mouthparts for capturing and manipulating food, sensory structures, or organs for transferring sperm. 4. The nervous system, which resembles that of the annelids, consists of a “brain” (cerebral ganglia) and ventral nerve cord with ganglia. In some arthropods, successive ganglia may fuse. Arthropods have a variety of very effective sense organs. Many have organs of hearing and antennae that sense taste and touch. Most insects and many crustaceans have compound eyes composed of many light-sensitive units called ommatidia (see Fig. 43-17). The compound eye can form an image and is especially adapted for detecting movement. Arthropods have an open circulatory system. A dorsal, tubular heart pumps hemolymph into a dorsal artery, which may branch into smaller arteries. From the arteries, hemolymph flows into large spaces that collectively make up the hemocoel. Eventually, hemolymph re-enters the heart through openings, called ostia, in its walls. The exoskeleton presents a barrier to diffusion of oxygen and carbon dioxide through the body wall, necessitating the evolution of specialized respiratory systems for gas exchange. Most aquatic arthropods have gills that function in gas exchange, whereas many terrestrial forms have a system of internal branching air tubes called tracheae, or tracheal tubes (see Fig. 46-1b). Other terrestrial arthropods have platelike book lungs (see Fig. 46-1d). The arthropod digestive system is a tube similar to that of the earthworm. Excretory structures vary somewhat among groups. The coelom is small and filled chiefly by the organs of the reproductive system. Arthropod evolution and classification are controversial Two groups of animals thought to be closely related to arthropods are the onychophorans and the tardigrades (FIG. 31-25). These three groups make up clade Panarthropoda. Like arthropods, onychophorans and tardigrades have a thick cuticle and must molt in order to grow. The body is segmented in all three groups, and all have legs and claws, and a ventral nervous system. Panarthropods have an open circulatory system, and the coelom is reduced to a hemocoel. The caterpillar-like onychophorans, known as velvet worms, probably branched early from the arthropod line. They have paired appendages, but they are not jointed. As in arthropods, their jaws are derived from appendages. Some biologists place the tardigrades, known as “water bears,” as the sister taxon of the arthropods. These tiny animals (typically less than 1 mm in length) have short, unjointed, clawed legs (see Fig. 2-12). Arthropod fossils have been identified dating back to the Proterozoic eon. Early in their evolutionary history, more than 540

Andrew Syred / Photo Researchers, Inc.

Dr. Morley Read / Photo Researchers, Inc.

view them as subphyla or classes. In this edition, we classify these groups as subphyla of phylum Arthropoda. However, as you study the following sections, bear in mind that arthropod systematics is a work in progress. Both the relationship of arthropods to other protostomes and the relationships among arthropods continue to be topics of lively debate.

Trilobites were early arthropods Among the earliest arthropods to evolve, trilobites inhabited shallow Paleozoic seas more than 500 mya. (a) Onychophoran. Note the soft (b) Tardigrades. Commonly known as Th ese arthropods, which have been extinct for about 250 segmented body and series of nonwater bears, tardigrades normally live jointed legs. Extant onychophorans in moist habitats, such as thin films of million years, lived on the sea bottom and filtered mud live in humid forests but many extinct water on mosses. Most are less than to obtain food. Most ranged from 3 to 10 cm (about 1 to species inhabited the ocean. 1 mm long. 4 in), but a few reached almost 1 m (39 in) in length. Covered by a hard, segmented exoskeleton, the FIGURE 31-25 Onychophorans and tardigrades trilobite body was a flattened oval divided into three These animals are thought to be close relatives of arthropods. parts: an anterior head bearing a pair of antennae and a pair of compound eyes; a thorax; and a posterior abmillion years ago (mya), arthropods diverged into several groups. domen (FIG. 31-26). At right angles to these divisions, two dorsal grooves extended the length of the animal, dividing the body into Arthropods evolved rapidly during the Cambrian radiation, as a median lobe and two lateral lobes. (The name trilobite derives evidenced by many diverse fossils in the Burgess Shale and other from this division of the body into three longitudinal parts.) Each ancient sites. segment had a pair of segmented biramous appendages, appendWe discuss five main arthropod groups, based on emerging ages with two jointed branches extending from their base. In trimolecular and other data: the extinct trilobites, and the extant lobites, each appendage consisted of an inner walking leg and an Myriapoda, Chelicerata, Crustacea, and Hexapoda (TABLE 31-5). Some systematists consider each of these groups a phylum; others outer branch with gills.

TABLE 31-5

Extant Arthropod Subphyla

Subphylum and Selected Classes

Body Divisions

Myriapoda Class Chilopoda (centipedes) Class Diplopoda (millipedes)

Head with segmented body

Chelicerata Class Merostoma (horseshoe crabs) Class Arachnida (spiders, scorpions, ticks, mites) Crustacea Class Malacostraca (lobsters, crabs, shrimp, isopods) Class Cirripedia* (barnacles) Class Copepoda* (copepods) Hexapoda Class Insecta (bees, ants, grasshoppers, roaches, flies, beetles)

Cephalothorax and abdomen

Head, thorax, and abdomen

Head, thorax, and abdomen

Appendages: Antennae, Mouthparts Uniramous. Antennae: one pair Mouthparts: mandibles, maxillae Uniramous. Antennae: none Mouthparts: chelicerae, pedipalps Biramous. Antennae: two pairs Mouthparts: mandibles, two pairs of maxillae (for handling food) Uniramous. Antennae: one pair Mouthparts: mandibles, maxillae

Appendages: Legs

Gas Exchange

Development

Main Habitat

Chilopods: one pair/segment Diplopods: usually two pairs/ segment

Tracheae

Direct

Terrestrial

Merostomes: four pairs of walking legs Arachnids: four pairs on cephalothorax Typically one pair/segment

Merostomes: gills Arachnids: book lungs or tracheae

Direct, except mites and ticks

Merostomes: marine Arachnids: mainly terrestrial

Gills

Usually larval stages (nauplius larva)

Marine or freshwater; a few are terrestrial

Three pairs on thorax

Tracheae

Typically, larval stages; most with complete metamorphosis

Mainly terrestrial

*Some systematists now include Cirripedia and Copepoda as orders under Class Maxillopoda.

Deuterostomia

Arthropods

Nematodes

Lophotrochozoans

Ctenophores

Cnidarians

Sponges

Ecdysozoa

Median lobe

William E. Ferguson

Lateral lobe

(a) Fossil trilobites. These extinct arthropods (Phacops rana) were about 3 cm (1.2 in) long. Note the large, well-developed eyes visible on either side of the head region. Antenna

Choanoflagellate ancestor

legs to merit their name; typically, they have approximately 30, although a few species have 100 or more. Because centipedes have long legs, they are able to run rapidly. Centipedes are carnivorous and feed on other animals, mostly insects. Larger centipedes eat snakes, mice, and frogs. Using their poison claws located just behind the head on the first trunk segment, centipedes capture and kill their prey. Millipedes (“thousand-legged”) have two pairs of legs on most body segments. Diplopods are not as agile as chilopods, and most species can crawl only slowly over the ground. However, they can powerfully force their way through earth and rotting wood. Millipedes are generally herbivorous and feed on both living and dead vegetation.

Head

Eye Gill

Lateral lobe Median lobe

(b) Dorsal view of a trilobite.

Chelicerates have no antennae Subphylum Chelicerata includes the merostomes (horseshoe crabs) and the arachnids. The chelicerate body consists of a cephalothorax (fused head and thorax) and an abdomen. Chelicerates are the only arthropods without antennae. They have no chewing mandibles. Instead, the first appendages, located immediately anterior to the mouth, are a pair of chelicerae (sing., chelicera), fanglike feeding appendages. Most chelicerates suck fluid food from their prey. The second appendages are the pedipalps. The chelicerae and pedipalps are modified to perform different functions in various groups, including manipulation of food, locomotion, defense, and copulation. The four pairs of legs on the cephalothorax are specialized for walking. Almost all the merostomes are extinct. Only the horseshoe crabs have survived, essentially unchanged for more than 350 million years. Limulus polyphemus, the species common along the Atlantic shore of North America, is horseshoe-shaped (FIG. 31-28a). Its long, spikelike tail is used in locomotion, not for defense or offense. Horseshoe crabs feed at night, capturing mollusks, worms, and other invertebrates that they find on the sandy ocean floor. Arachnids include spiders, scorpions, ticks, harvestmen (daddy longlegs), and mites (FIGS. 31-28b and c). Most of the approximately 80,000 named species are carnivorous and prey on insects and other small arthropods. Arachnids have six pairs of jointed appendages. In spiders, the first appendages, the chelicerae, are used to penetrate prey; some species use the chelicerae to inject poison into their prey. Spiders use the second pair of appendages, the pedipalps, to hold and chew food. Many arachnid species have pedipalps modified as sense organs for tasting food or for repro-

Biramous appendages

(c) Ventral view.

FIGURE 31-26 Trilobites Biologists consider these extinct marine arthropods the most primitive members of phylum Arthropoda. Trilobites flourished in the ocean during the Paleozoic era.

Subphylum Myriapoda includes centipedes and millipedes Members of subphylum Myriapoda are characterized by unbranched appendages called uniramous appendages, jawlike mandibles, and a single pair of antennae. Centipedes (class Chilopoda) and millipedes (class Diplopoda) are terrestrial and are typically found beneath stones or wood in the soil in both temperate and tropical regions. Both centipedes and millipedes have a head and an elongated trunk with many segments, each bearing uniramous legs (FIG. 31-27). Centipedes (“hundred-legged”) have one pair of legs on each segment behind the head. Most centipedes do not have enough

Dwight Kuhn

Milton H. Tierney Jr. / Visuals Unlimited

(a) Horseshoe crabs.The only living merostomes are a few closely related species of horseshoe crabs. Seasonally, horseshoe crabs return to beaches for mating. In this photograph Limulus polyphemus males are competing for a female.

Steve Maslowski/Visuals Unlimited

.John R. MacGregor / Peter Arnold, Inc.

(a) Centipede (Lithobius ), a member of class Chilopoda. Centipedes have one pair of uniramous appendages per segment.

(b) Millipede (Diplopoda pachydesmus), a member of class Diplopoda. Millipedes have two pairs of uniramous appendages per segment.

duction (for sperm transfer and courtship displays). Scorpions use their very large pedipalps as pincers for capturing prey. Chelicerates use the remaining four pairs of appendages for walking. Typically, spiders have eight eyes arranged in two rows of four each along the anterior dorsal edge of the cephalothorax. The eyes detect movement, locate objects, and in some species form a relatively sharp image. Gas exchange in arachnids takes place by either tracheal tubes, book lungs, or both. A book lung consists of 15 to 20 parallel plates (like pages of a book) that contain tiny blood vessels. Air enters the body through abdominal slits and circulates between the plates. As air passes over the blood vessels in the plates, oxygen diffuses into the blood and carbon dioxide diffuses out of the blood into the air. As many as four pairs of book lungs provide an extensive surface area for gas exchange. The spider has unique glands in its abdomen that secrete silk, an elastic protein that is spun into fibers by organs called spinnerets.

▲

FIGURE 31-28 Chelicerates

K. H. Kjeldsen/Science Photo Library/Photo Researchers, Inc.

(b) A female black widow spider (Latrodectus mactans ) rests on her web. The venom of the black widow spider is a neurotoxin.

FIGURE 31-27 Myriapods

100 μm

(c) The house dust mite Dermatophagoides. This mite, a common inhabitant of homes, is associated with house dust allergies (color enhanced SEM).

(a) Goose barnacles (Lepas sp.) filter feeding. These stalked barnacles are found in large numbers on intertidal rocks. The ones in this image are attached to a fisherman’s float, North Pacific.

© Biosphoto/ Danna Patricia

Crustaceans are vital members of marine food webs Subphylum Crustacea includes lobsters, crabs, shrimp, barnacles, and their relatives (FIG. 31-29). Many crustaceans are primary consumers of algae and detritus. Countless billions of microscopic crustaceans are part of marine zooplankton, the free-floating, mainly microscopic organisms in the upper layers of the ocean. These crustaceans are food for many fishes and other marine animals, such as certain baleen whales. Paleobiologists have dated crustacean fossils back to the Cambrian period (more than 505 mya). A distinctive feature of crustaceans is the nauplius larva, which is often the first stage after hatching. This larva has only the most anterior three pairs of appendages. Crustaceans are also characterized by mandibles, biramous appendages, and two pairs of antennae. Their antennae, which are the first and second pairs of appendages, serve as sensory organs for touch and taste. Most adult crustaceans have compound eyes, and many crustaceans have statocysts, sense organs that detect gravity (see Fig. 43-6). The hard mandibles, used for biting and grinding food, are the third pair of appendages; they lie on each side of the ventral mouth. Posterior to the mandibles are two pairs of appendages, the first and second maxillae, used for manipulating and holding food. Several other pairs of appendages are present. Some are modified for walking. Others may be specialized for swimming, sperm transmission, carrying eggs and young, or sensation.

Norbert Wu / Minden Pictures/ NGS Image Collection

The silk is liquid as it emerges from the spinnerets but hardens after it leaves the body. Spiders use silk to build nests, to encase their eggs in a cocoon, and in some species, to trap prey in a web. Many spiders lay down a silken dragline that serves both as a safety line and a means of communication between members of a species. From the type of dragline, another spider can determine the sex and maturity level of the spinner. Although all spiders (and scorpions) have poison glands useful in capturing prey, only a few produce poison toxic to humans. The most widely distributed poisonous spider in the United States is the black widow (see Fig. 31-28b). Its venom is a neurotoxin that interferes with transmission of messages from nerves to muscles. The shiny black female is about 3.8 cm (1.5 in) long with a red or orange hourglass-shaped marking on the underside of its abdomen. Adult male black widow spiders are smaller and are harmless. After mating, the female sometimes kills and eats the male. The brown recluse spider (Loxosceles reclusa) is more slender than the black widow and has a violin-shaped stripe on its dorsal cephalothorax. Its venom destroys the tissues surrounding the bite. Although painful, spider bites cause fewer than three fatalities per year in the United States. Although some mites contribute to soil fertility by breaking down organic matter, many are serious nuisances. They eat crops, infest livestock and pets, and inhabit our own bodies. Certain mites cause mange in dogs and other domestic animals. Chiggers (red bugs), the larval form of red mites, attach themselves to the skin and secrete an irritating digestive fluid that may cause itchy red welts. Larger than mites, ticks are parasites on dogs, deer, and many other animals. They transmit the bacteria that cause diseases such as Rocky Mountain spotted fever and Lyme disease (see Fig. 54-2).

(b) Sponge crab (Criptodromia octodenta) wearing a sponge on its back. The sponge provides camouflage and shelter. The sponge crab uses its claws to cut out a fragment from a sponge and then trims the sponge to conform to its own shape. The last two pairs of the crab’s legs bend upward over the crab’s carapace and hold the sponge in place. The sponge grows along with the crab.

FIGURE 31-29 Crustaceans

Crustaceans are the only group of arthropods that are primarily aquatic; not surprisingly, they typically have gills for gas exchange. Two large glands located in the head excrete metabolic wastes and regulate salt balance. Most crustaceans have separate sexes. During copulation, the male uses specialized appendages to transfer sperm

Insects are articulated, tracheated hexapods With more than one million described species (and perhaps millions more not yet identified), class Insecta of subphylum

Cephalothorax Abdomen

Thorax

Head

Photo by Tim Dyes, Grand Cayman

into the female. Newly hatched animals may resemble adults, or they may undergo successive molts as they pass through a series of larval stages before developing an adult body. Barnacles, the only sessile crustaceans, differ markedly in external anatomy from other members of the subphylum. They are marine suspension feeders that secrete limestone cups within which they live (see Fig. 31-29a). The larvae of barnacles are free-swimming forms that go through several molts. They eventually become sessile and develop into the adult form. The 19th-century naturalist Louis Agassiz described the barnacle as “nothing more than a little shrimplike animal standing on its head in a limestone house and kicking food into its mouth.” The bane of marine boaters, barnacles can proliferate on ship bottoms in great numbers. Barnacles can reduce the speed of a ship by more than 10% and increase fuel consumption by as much as 40%. The United States Office of Naval Research has been testing an underwater robot that cleans barnacles and other marine life from the hulls of ships when they are in port. Isopods are mainly tiny (5 to 15 mm in length) marine crustaceans that inhabit the ocean floor. However, this group also includes some terrestrial animals: pill bugs and sow bugs. The most abundant crustaceans are the copepods, which are typically microscopic. Marine copepods are the most numerous component of zooplankton. The largest and most familiar order of crustaceans, Decapoda, contains more than 10,000 species of lobsters, crayfish, crabs, and shrimp. Most decapods are marine, but a few, such as the crayfish, certain shrimp, and a few crabs, live in fresh water. The crustaceans in general and the decapods in particular show striking specialization and differentiation of parts in the various regions of the animal. In the lobster, the appendages in the different parts of the body differ markedly in form and function (FIG. 31-30). The five segments of the lobster’s head and the eight segments of its thorax are fused into a cephalothorax, which is covered on the top and sides by a shield, the carapace, composed of chitin impregnated with calcium salts. The two pairs of antennae serve as chemoreceptors and tactile sense organs. The mandibles are short and heavy, with opposing surfaces used in grinding and biting food. Behind the mandibles are two pairs of accessory feeding appendages: the first and second maxillae. The appendages of the first three segments of the thorax are the maxillipeds, which aid in chopping up food and passing it to the mouth. The fourth segment of the thorax has a pair of large chelipeds, or pinching claws. The last four thoracic segments bear walking legs. The appendages of the first abdominal segment are part of the reproductive system and are used by the male to transfer sperm. The following four abdominal segments bear paired swimmerets, small paddlelike structures used by some decapods for swimming and by the females of all species for holding eggs. Each branch of the sixth abdominal appendages consists of a large flattened structure. Together with the flattened posterior end of the abdomen, they form a tail fan used for swimming backward.

(a) Tail fan

Fifth walking leg

Eye Second Third walking maxilliped leg Cheliped Mouth

First antenna Second antenna

Swimmerets First swimmeret (used by male (b) in copulation)

FIGURE 31-30 Anatomy of the lobster (a) Like other decapods, the spiny lobster (Panulirus argus) has five pairs of walking legs. The first pair of walking legs is modified as chelipeds (large claws). Photographed in Grand Cayman. (b) Ventral view of a lobster. Note the variety of specialized appendages.

Hexapoda is the most successful group of animals on our planet in terms of number of individuals, number of species, diversity, and geographic distribution (TABLE 31-6). What they lack in size, insects make up in sheer numbers. If we could weigh all the insects in the world, their weight would exceed that of all the remaining terrestrial animals. Although primarily terrestrial, some insect species live in fresh water, a few are truly marine, and others inhabit the shore between the tides. The earliest fossil insects—primitive, wingless species—date to the Devonian period more than 360 mya. Insect fossils from the Carboniferous period more than 310 mya include both wingless

TABLE 31-6

Some Orders of Insects

Order, Number of Species, Some Characteristics Homoptera (43,000)* Leafhoppers, scale insects Two pairs of membranous wings; piercing–sucking mouthparts form beak; parasites of plants; vectors of plant diseases; incomplete metamorphosis Odonata (6000) Dragonflies, damselflies Two pairs of long membranous wings; chewing mouthparts; large, compound eyes; active predators; larvae very different from adult; incomplete metamorphosis Orthoptera (20,000) Grasshoppers, crickets Forewings leathery, hindwings membranous; chewing mouthparts; most herbivorous, some cause crop damage; some predatory; incomplete metamorphosis Blattodea (4000) Cockroaches When wings present, forewings leathery, hindwings membranous; chewing mouthparts; legs adapted for running; incomplete metamorphosis

Representative Adult

Buffalo treehopper Stictocephala bubalus

Hemiptera (80,000) Chinch bugs, bedbugs, water striders; cicadas and aphids are now assigned to this order Hindwings membranous; forewings smaller; piercing–sucking mouthparts form beak; most herbivorous; some parasitic; incomplete metamorphosis

Representative Adult

Lepidoptera (160,000) Moths, butterflies Usually two pairs of membranous, colorful, scaled wings; larvae are wormlike caterpillars with chewing mouthparts for eating plants; adult mouthparts adapted for sucking nectar Luna moth Aetias luna

Fork-tailed bush katydid Scudderia furcata

Diptera (150,000) Houseflies, mosquitoes, fruit flies Only forewings functional in flying; hindwings reduced to small, knoblike balancing organs (halteres); mouthparts usually adapted for sucking (and piercing in some); larvae are maggots that damage domestic animals or food; adults transmit diseases such as sleeping sickness, yellow fever, and malaria

American cockroach Periplaneta americana

Siphonaptera (2000) Fleas No wings; piercing–sucking mouthparts; legs adapted for clinging and jumping; parasites on birds and mammals; vectors of bubonic plague and typhus

Dog flea Ctenocephalides canis

Eastern subterranean termite Reticulitermes flavipes

Coleoptera (360,000) Beetles, weevils Forewings modified as protective coverings for membranous hindwings (which are sometimes absent); chewing mouthparts; largest order of insects; most herbivorous; some aquatic

Colorado potato beetle Leptinotarsa decemlineata

Damselfly Ischnura

Isoptera (2100) Termites Two pairs of wings, or none; wings shed by sexual forms after mating; chewing mouthparts; social insects, form large colonies; eat wood; incomplete metamorphosis

Phthiraptera (3000) Lice, sucking lice No wings; piercing–sucking mouthparts; ectoparasites of mammals; head and body louse and crab louse are human parasites; vectors of typhus fever; incomplete metamorphosis

Order, Number of Species, Some Characteristics

Human head and body louse Pediculus humanus

Chinch bug Blissus leucopterus

*The number of species listed for each order is approximate. Estimates vary widely.

Hymenoptera (190,000) Ants, bees, wasps Usually two pairs of membranous wings; mouthparts may be modified for sucking or lapping nectar; many are social insects; some sting; important pollinators

Thysanura (320) Silverfish No wings; biting–chewing mouthparts; two to three “tails” extend from posterior tip of abdomen; inhabit dead leaves; eat starch in books; direct development

Deerfly Chrysops vittatus

Bald-faced hornet Vespula maculata

Silverfish Lepisma saccharina

and primitive winged species. Cockroaches, mayflies, and cicadas are among the insects that have survived relatively unchanged from the Carboniferous period to the present day. Molecular and developmental studies suggest that hexapods are not a monophyletic group and that they are closely related to the crustaceans. At the time of this writing, many biologists classify crustaceans along with hexapods in clade Pancrustacea. Some systematists hypothesize that crustaceans and hexapods are sister taxa, closely related taxa that occupy adjacent branches in a cladogram (see Chapter 23). Others argue that hexapods evolved within the crustacean clade. We describe an insect as an articulated (jointed), tracheated (having tracheal tubes for gas exchange) hexapod (having six feet). The insect body consists of three distinct parts: head, thorax, and abdomen (FIG. 31-31). An insect’s uniramous (unbranched) appendages include three pairs of legs that extend from the adult thorax and, in many orders, one or two pairs of wings. One pair of antennae protrudes from the head, and the sense organs include both simple and compound eyes. The complex mouthparts, which include mandibles and maxillae, are adapted for piercing, chewing, sucking, or lapping. The tracheal system of insects has made an important contribution to their diversity. Air enters the tracheal tubes, or tracheae, through spiracles, tiny openings in the body wall. Oxygen passes directly to the internal organs. This effective oxygen delivery system permits insects to have the high metabolic rate necessary for activities such as flight, even though they have an open circulatory system. Excretion is accomplished by two or more Malpighian tubules, which receive metabolic wastes from the blood, concentrate the wastes, and discharge them into the intestine. Unique to terrestrial arthropods, Malpighian tubules also perform the extremely important function of conserving water (see Fig. 48-4). The sexes are separate, and fertilization takes place internally. Several molts occur during development. The immature stages between molts are called instars. Primitive insects with no wings, such as silverfish, have direct, or simple, development: the young hatch as juveniles that resemble the adult form. Other insects, such as grasshoppers and cockroaches, undergo incomplete metamorphosis in which the egg gives rise to a larva that resembles the adult in many ways but lacks functional wings and reproductive structures. The larva goes through a series of molts during which it becomes more and more like the adult. Most insects, including bees, butterflies, and fleas, undergo complete metamorphosis with four distinct stages in the life cycle: egg, larva, pupa, and adult (FIG. 31-32). The wormlike larva does not look at all like the adult. For example, caterpillars have a body form entirely different from that of butterflies. Typically, an insect spends most of its life as a larva, which has chewing mouthparts. Eventually, the larva stops feeding, molts, and enters a pupal stage, usually within a protective cocoon or underground burrow. The pupa does not feed and typically cannot defend itself. Energy reserves stored during the larval stage are spent remodeling its body. When it emerges as an adult, it is equipped with functional wings and reproductive organs.

KEY POINT

Insects, the most successful of all animals, are articulated, tracheated hexapods.

Head

Thorax Abdomen

Forewing

Antenna Simple eye Compound eye Sound receptor

Spiracles

Hindwing

(a) External structure. Note the three pairs of segmented legs.

Ovary Digestive gland Heart

Anus

Brain

Nerve cord Intestine Malpighian tubules Genital opening

(b) Internal anatomy.

FIGURE 31-31 Animated Anatomy of the grasshopper

Certain species of bees, ants, and termites exist as colonies or societies made up of several different types of individuals, each adapted for some particular function. The members of some insect societies communicate with one another by “dances” and by means of pheromones, substances secreted to the external environment. Social insects and their communication are discussed in Chapter 52. Adaptations that contribute to the biological success of insects

What are the secrets of insect success? The tough exoskeleton protects insects from predators and helps reduce water loss. Segmentation allows mobility and flexibility and permits regional specialization of the body. For example, cephalization and highly developed sense organs are important in both offense and defense. The insect body plan has been modified and specialized in so many ways that insects are adapted to a remarkable number of lifestyles. The jointed appendages are specialized for various types of feeding, sensory functions, and different types of locomotion (walking, jumping, swimming). For example, grasshopper mouthparts are adapted for biting and chewing leaves. Moth and butterfly mouthparts are adapted for sucking nectar from flowers, and mosquito mouthparts are adapted for sucking blood. Aphids and leafhoppers have mouthparts specialized to pierce plants and feed on plant juices. Another adaptation that contributes to insect success is the ability to fly. Unlike other terrestrial invertebrates, which creep slowly along on or under the ground, many insects fly rapidly through the air. Their wings and small size facilitate their wide distribution and,

KEY POINT

Metamorphosis reduces competition among insects within the same species; larval forms do not compete with adults for food or habitats.

© Millard H. Sharp / Photo Researchers, Inc.

James H. Robinson / Photo Researchers, Inc.

Matt Meadows / Peter Arnold, Inc.

(b) Caterpillar feeding on sweet gum leaf. Adam Jones / Photo Researchers, Inc.

(a) Polyphemus moth caterpillars hatching from eggs.

(c) Cocoon cut open to show developing pupa.

(d) Adult on tree. Note eyespots.

FIGURE 31-32 Animated Complete metamorphosis in a Polyphemus moth This moth (Antheraea polyphemus) belongs to the family of giant silk moths.

in many instances, their immediate survival. For example, when a pond dries up, adult aquatic insects can fly to another habitat. The reproductive capacity of insects is remarkable. Under ideal conditions, the fruit fly Drosophila can produce 25 generations in a single year! Insect eggs are protected by a thick membrane, and several eggs may, in addition, be enclosed in a protective egg case. By dividing the insect life cycle into different stages, metamorphosis reduces competition among members of the same species (intraspecific competition). For example, larval forms have different lifestyles, so they do not compete with adults for food or habitats. Insects have many interesting adaptations for offense and defense (see Fig. 7-8). We all are familiar with the stingers of bees and wasps, which are specialized egg-laying structures (ovipositors). Many insects have cryptic coloration, which allows them to blend into the background of their habitat; some look like dead twigs or leaves. Others mimic poisonous insects, deriving protection from this resemblance. Still others “play dead” by remaining motionless. Insects communicate by tactile, auditory, visual, or chemical signals. For example, certain ant species use pheromones to mark trails and to warn of danger. Other insects use pheromones to attract a mate.

Impact of insects on humans Not all insects compete with humans for food or cause us to scratch, swell up, or recoil from their presence. Bees, wasps, beetles, and many other insects pollinate the flowers of crops and fruit trees. Some insects destroy other insects that are harmful to humans. For example, dragonflies eat mosquitoes. And many organic farmers and home gardeners buy ladybird beetles, which are adept at ridding plants of aphids and other insect pests. Insects are important members of many food webs. Many birds, mammals, amphibians, reptiles, and some fishes depend on insects for food. Some beetles and the larvae (maggots) of flies are detritus feeders—they break down dead plants and animals and their wastes, permitting nutrients to be recycled. Various insect products are useful to humans. Bees produce honey as well as beeswax, which we use to make candles, lubricants, and other products. Shellac is made from lac, a substance given off by certain scale insects that feed on the sap of trees. And the labor of silkworms provides us with beautiful fabric. On the negative side, insects destroy crops worth billions of dollars each year. Termites destroy buildings, and moths damage clothing. Fire ants not only inflict painful stings but cause farmers serious economic loss because of their large mounds, which damage mowers and other farm equipment. Blood-sucking flies, screwworms, lice, fleas, and other insects annoy and transmit dis-

ease to humans and domestic animals. Mosquitoes are vectors of yellow fever and malaria (see Fig. 26-7). Fleas and lice transmit the rickettsia that cause typhus, and houseflies sometimes transmit typhoid fever and dysentery. Tsetse flies transmit African sleeping sickness, and fleas may be vectors of bubonic plague. In this chapter, we have surveyed the major groups of protostomes. TABLE 31-7 summarizes these groups. Chapter 32 focuses on the deuterostomes, which include the echinoderms and chordates.

TABLE 31-7

Review ■ ■

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What characters distinguish nematodes from flatworms? What are four key arthropod characters, and how does each contribute to arthropod success? How do myriapods differ from chelicerates? What are the functions of each of these appendages in a lobster: maxillae, chelipeds, and swimmerets? Define an insect. What are four adaptations that have contributed to insect success?

Overview of Sponges, Cnidarians, Ctenophores, and Protostomes

Major Groups

Body Plan

Some Key Characteristics

Poriferans (sponges)

Asymmetrical; cells loosely arranged and do not form true tissues; body is sac with pores, central cavity, and osculum

Collar cells (choanocytes); aquatic, mainly marine

Cnidarians (hydras, jellyfish, corals)

Radial symmetry; diploblastic; gastrovascular cavity with one opening

Tentacles with cnidocytes (stinging cells) that discharge nematocysts; two body forms: polyp and medusa; some form colonies; mainly marine

Ctenophores (comb jellies)

Biradial symmetry; diploblastic; gastrovascular cavity with mouth and anal pores

Eight rows of cilia that resemble combs; tentacles with adhesive glue cells; marine predators

Flatworms (planarians, tapeworms, flukes)

Bilateral symmetry; triploblastic; simple organ systems; gastrovascular cavity with one opening

No body cavity; some cephalization; some are free-living carnivores; many are parasites

Nemerteans (ribbon worms)

Bilateral symmetry; triploblastic; organ systems; complete digestive tube*

Proboscis (long, muscular tube that can be everted to capture prey); true coelom reduced; mainly marine carnivores

Mollusks (clams, snails, squids)

Bilateral symmetry; triploblastic; organ systems; complete digestive tube; true coelom

Soft body usually covered by dorsal shell; flat, muscular foot; mantle covers visceral mass; most have a radula (belt of teeth)

Annelids (some marine worms, earthworms, leeches)

Bilateral symmetry; triploblastic; organ systems; complete digestive tube; true coelom

Segmented body; most have setae, bristles that provide traction during crawling

Lophophorates (brachiopods, phoronids, bryozoans)

Bilateral symmetry; triploblastic; organ systems; complete digestive tube; true coelom

Lophophore (ciliated ring of tentacles) surrounds mouth, captures suspended particles in water; mainly sessile, marine animals

Rotifers (wheel animals)

Bilateral symmetry; triploblastic; organ systems; complete digestive tube; pseudocoelom

Crown of cilia at anterior end; cell number constant; microscopic, aquatic animals

Nematodes (roundworms, e.g., Ascaris, hookworms, trichina worms)

Bilateral symmetry; triploblastic; organ systems; complete digestive tube; pseudocoelom

Cylindrical, threadlike body; widely distributed in soil and aquatic sediments; important as decomposers; many are predators; some are parasites

Arthropods (centipedes, spiders, crabs, lobsters, insects)

Bilateral symmetry; triploblastic; organ systems; complete digestive tube; true coelom

Segmented body; exoskeleton; paired, jointed appendages; insects and many crustaceans have compound eyes; insects have tracheal tubes for gas exchange

Protostomes: Lophotrochozoan Branch

Protostomes: Ecdysozoan Branch

*A complete digestive tube has a mouth for food intake and an opening (anus) for elimination of wastes.

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

Deuterostomia

Ecdysozoans

Rotifers

Lophophorates

Annelids

Mollusks

Nemerteans

Platyhelminthes

The poriferans, or sponges, are characterized by flagellate collar cells (choanocytes) that generate a water current that brings food and oxygen to the cells. Collar cells also trap and phagocytize food particles. The sponge body is a sac with tiny openings through which water enters; a central cavity, or spongocoel; and an open end, or osculum, through which water exits. The cells of sponges are loosely associated; they do not form true tissues.

Sponges

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The Lophotrochozoa make up a clade that includes some of the flatworms, nemerteans, mollusks, annelids, the lophophorate phyla, and rotifers.

Cniderians

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1 Identify distinguishing characteristics of poriferans.

Ctenophores

31.1 (page 641)

Learn more about sponge structure by clicking on the figure in CengageNOW.

2 Identify distinguishing characteristics of cnidarians, describe four groups, and give examples of animals that belong to each group. Cnidarians are characterized by radial symmetry, two tissue layers, and cnidocytes, cells containing stinging organelles called nematocysts. The gastrovascular cavity has a single opening that serves as both mouth and anus. Nerve cells form irregular, nondirectional nerve nets that connect sensory cells with contractile and gland cells. The life cycle of many cnidarians includes a sessile polyp stage ■ (a form with a dorsal mouth surrounded by tentacles) and a freeswimming medusa (jellyfish) stage. ■ Phylum Cnidaria includes four groups. Hydrozoa (hydras, hydroids, and the Portuguese man-of-war) are typically polyps and may be solitary or colonial. Scyphozoa (jellyfish) are generally medusae. Cubozoa, the “box jellyfish,” have complex eyes that form blurred images. Anthozoa (sea anemones and corals) are polyps and may be solitary or colonial; anthozoans differ from hydrozoans in the organization of the gastrovascular cavity. ■

Learn more about cnidarian body forms, nematocysts, and life cycles by clicking on the figures in CengageNOW.

3 Identify distinguishing characteristics of ctenophores. ■

Ctenophores, or comb jellies, are fragile, luminescent marine predators with biradial symmetry. Ctenophores have eight rows of cilia that resemble combs. They are diploblastic and have tentacles with adhesive glue cells.

31.2 (page 648) 4 Summarize what is known about the phylogeny of the lophotrochozoans and cite the adaptive advantages of having a coelom and of cephalization.

Choanoflagellate ancestor

The true coelom is a fluid-filled body cavity completely lined by mesoderm that lies between the digestive tube and the outer body wall. The coelom brings about the tube-within-a-tube body plan. The body wall is the outer tube. The inner tube is the digestive tube. The coelom can serve as a hydrostatic skeleton in which contracting muscles push against a tube of fluid. The coelom is a space in which internal organs, including gonads, can develop; it helps transport materials; and protects internal organs. ■ Cephalization, the evolution of a head with the concentration of sense organs and nerve cells at the anterior end, increases the effectiveness of a bilateral animal to actively find food, shelter, and mates and to detect enemies. 5 Identify distinguishing characteristics of flatworms and nemerteans; describe the main groups of flatworms, giving examples of animals that belong to each class. ■ The flatworms are acoelomate (have no coelom) animals with bilateral symmetry, cephalization, three definite tissue layers, and well-developed organs. Many flatworms are hermaphrodites; a single animal produces both sperm and eggs. Flatworms have a ladder-type nervous system, typically consisting of sense organs and a simple brain composed of two ganglia. The ganglia are connected to two nerve cords that extend the length of the body. Protonephridia function in osmoregulation and disposal of metabolic wastes. ■ Four groups of flatworms are recognized: class Turbellaria comprises free-living flatworms, including planarians; classes Trematoda and Monogenea include the parasitic flukes; class Cestoda includes the parasitic tapeworms. The parasitic flukes and tapeworms typically have suckers or hooks for holding on to their hosts; they have complicated life cycles with intermediate hosts and produce large numbers of eggs. ■

Watch the fluke and tapeworm life cycles by clicking on the figures in CengageNOW. ■

Nemerteans (ribbon worms) are characterized by the proboscis, a muscular tube used in capturing food and in defense. Nemerteans have a tube-within-a-tube body plan, a complete digestive tract with mouth and anus, and a circulatory system. The coelom is reduced.

6 Describe the distinguishing characteristics of mollusks and of the four main groups of mollusks discussed, giving examples of each group. ■ Mollusks are soft-bodied animals usually covered by a shell. They have a ventral foot for locomotion and a pair of folds called the mantle that covers the visceral mass, a concentration of body organs. ■ Mollusks have an open circulatory system except for cephalopods, which have a closed circulatory system. A rasplike radula functions as a scraper in feeding in all groups except the bivalves, which are filter feeders. Typically, marine mollusks have a free-swimming, ciliated trochophore larva. ■ Polyplacophorans are chitons, mollusks with shells consisting of eight overlapping dorsal plates. The gastropods, which include the snails, slugs, and their relatives, have a well-developed head with tentacles. The body undergoes torsion, a twisting of the visceral mass. Bivalves are aquatic clams, scallops, and oysters. A two-part shell, hinged dorsally, encloses the bodies of these filter feeders. Cephalopods include the squids, octopods, and Nautilus. These active, predatory swimmers have tentacles surrounding the mouth, which is located in the large head. 7 Describe the distinguishing characteristics of annelids and of the three main groups of annelids discussed, giving examples of each group. ■ The annelids, the segmented worms, include many aquatic worms, earthworms, and leeches. Annelids have long bodies with segmentation both internally and externally; their large, compartmentalized coelom serves as a hydrostatic skeleton. ■ Polychaetes are marine annelids characterized by parapodia, appendages used for locomotion and gas exchange. The parapodia bear many bristlelike structures called setae. Polychaetes also differ from other annelids in having a well-defined head with sense organs. Oligochaetes, the group that includes the earthworms, are characterized by a few short setae per segment. The body is divided into more than 100 segments separated internally by septa. Leeches belong to the group Hirudinida. Setae and appendages are absent. Parasitic leeches are equipped with suckers for holding on to their host. 8 Describe the distinguishing characteristics of the lophophorates and of the rotifers. ■ The lophophorates, marine animals that have a lophophore, include the brachiopods, phoronids, and bryozoans. The lophophore, a ciliated ring of tentacles surrounding the mouth, is specialized for capturing suspended particles in the water. ■ Rotifers are pseudocoelomates that are thought to have evolved from animals with a true coelom. They have a crown of cilia at their anterior end.

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Phylum Arthropoda is composed of segmented animals with paired, jointed appendages and an armorlike exoskeleton of chitin. Molting is necessary for the arthropod to grow. Arthropods have an open circulatory system with a dorsal heart that pumps hemolymph. Aquatic forms have gills for gas exchange; terrestrial forms have either tracheae or book lungs. The arthropods along with the onychophorans (velvet worms) and tardigrades (water bears) make up the clade Panarthropoda. Based on molecular and other data, arthropods are currently assigned to five main groups: extinct trilobites and extant Myriapoda, Chelicerata, Crustacea, and Hexapoda. The trilobites are extinct marine arthropods covered by a hard, segmented shell. Each segment had a pair of biramous appendages, appendages with two jointed branches, an inner walking leg and an outer gill branch. Subphylum Myriapoda includes class Chilopoda, the centipedes, and class Diplopoda, the millipedes. Members of this subphylum have uniramous (unbranched) appendages and a single pair of antennae. Subphylum Chelicerata includes the merostomes (horseshoe crabs) and the arachnids (spiders, mites, and their relatives). The chelicerate body consists of a cephalothorax and abdomen; there are six pairs of uniramous, jointed appendages, of which four pairs serve as legs. The first appendages are chelicerae; the second are pedipalps. These appendages are adapted for manipulation of food, locomotion, defense, or copulation. Chelicerates have no antennae and no mandibles. Subphylum Crustacea includes lobsters, crabs, shrimp, pill bugs, and barnacles. The body consists of a cephalothorax and abdomen. Crustaceans typically have five pairs of walking legs. Appendages are biramous. Crustaceans have two pairs of antennae that sense taste and touch. The third appendages are mandibles used for chewing. Two pairs of maxillae, posterior to the mandibles, manipulate and hold food. Subphylum Hexapoda includes class Insecta. An insect is an articulated, tracheated hexapod; its body consists of head, thorax, and abdomen. Insects have uniramous appendages, a single pair of antennae, tracheae for gas exchange, and Malpighian tubules for excretion.

31.3 (page 661) 9 Summarize what is known about the phylogeny of the ecdysozoans. Ecdysozoa is one of the three major animal clades; its validity is based on many types of evidence, including molecular data. Members of this group go through the process of ecdysis, or molting, during which an animal sheds its outer covering, which is then replaced by the growth of a new one. 10 Describe distinguishing characteristics of nematodes and identify four parasitic nematodes. Nematodes, or roundworms, have a pseudocoelom. The body is ■ covered by a tough cuticle that helps prevent desiccation. Parasitic nematodes that infect humans include Ascaris, hookworms, trichina worms, and pinworms. 11 Describe six key characteristics of arthropods, summarize proposed phylogeny of arthropods, and distinguish among arthropod subphyla and classes; give examples of animals that belong to each group. ■

12 Identify adaptations that have contributed to the biological success of insects. ■ The biological success of the insects can be attributed to their many adaptations, including a versatile exoskeleton, segmentation, specialized jointed appendages, highly developed sense organs, and ability to fly. Metamorphosis, transition from one developmental form to another, reduces competition within the same species. Insects have developed effective reproductive strategies, effective mechanisms for defense and offense, and the ability to communicate. Explore the body plans of the protostomes by clicking on the figures in CengageNOW.

T E S T YO U R U N D E R S TA N D I N G 1. Collar cells (choanocytes) are characteristic of (a) poriferans (b) cnidarians (c) most coelomates (d) lophotrochozoans (e) ecdysozoans 2. Cnidocytes (a) are characteristic of sponges (b) are found among cells lining the gastrovascular cavity (c) contain stinging organelles (d) are lined with mesoderm (e) have two main shapes 3. Which of the following is associated with the evolution of the coelom? (a) radial symmetry (b) tube-within-a-tube body plan (c) incomplete metamorphosis (d) bilateral symmetry (e) development of ganglia 4. Trochophore larvae are characteristic of (a) arthropods (b) cnidarians (c) flatworms (d) mollusks (e) crustaceans 5. Which of the following is not an adaptation to parasitic life? (a) production of a few well-protected eggs (b) hooks (c) suckers (d) reduced digestive system (e) intermediate host 6. An open circulatory system (a) is characteristic of squids and other active mollusks (b) permits contracting muscles to recover quickly (c) has no blood vessels (d) is a unique characteristic of annelids (e) has hemolymph, which bathes tissues directly

arthropods (e) are an evolutionary link between annelids and arthropods 9. Which of the following is not characteristic of arthropods? (a) exoskeleton (b) pseudocoelom (c) paired, jointed appendages (d) chitin (e) segmentation 10. Which of the following is characteristic of insects? (a) biramous appendages (b) two pairs of antennae (c) chelicerae (d) eight legs (e) mandibles 11. The correct sequence of complete metamorphosis of an insect is (a) egg ¡ immature form ¡ adult (b) egg ¡ trochophore larva ¡ adult (c) egg ¡ pupa ¡ larva ¡ adult (d) egg ¡ larva ¡ pupa ¡ adult (e) adult ¡ larva ¡ egg ¡ pupa 12. Label the diagram. Refer to Figure 31-31 b to confirm your answers.

7. Which of the following characteristics is associated with mollusks? (a) mandibles (b) mantle (c) pedipalps (d) chelipeds (e) setae 8. Trilobites (a) were early mollusks (b) are onychophorans (c) are characterized by parapodia and setae (d) were early

CRITICAL THINKING

Deuterostomia

Ecdysozoans

Lophotrochozoans

Ctenophores

Cnidarians

Sponges

1. ANALYZING DATA. Imagine that you discover a new animal in a rain forest. It has an elongated, segmented body. Where would you place this animal in the cladogram below? Support your decision. Look at the more detailed cladogram in Figure 30-7a (in Chapter 30). What additional characteristics would help you place your animal in this cladogram? Explain your decision-making process.

?

Choanoflagellate ancestor

2. EVOLUTION LINK. Acoelomates, pseudocoelomates, and coelomates were historically classified as distinct groups of animals. In what way has this classification changed? Why have biologists stopped using type of body cavity as a major criterion for inferring relationships among animal groups?

3. EVOLUTION LINK. Animal body plans have generally evolved from simple to more complex. How can we explain the exceptions? 4. EVOLUTION LINK. Discuss important adaptations made possible by the development of the coelom. 5. EVOLUTION LINK. Discuss the idea that every evolutionary adaptation has both advantages and disadvantages, using each of the following as an example: (a) cephalization, (b) the arthropod exoskeleton, and (c) segmentation with specialization. 6. EVOLUTION LINK. Insects that undergo complete metamorphosis outnumber those that do not by more than 10 to 1. Hypothesize an explanation. 7. SCIENCE, TECHNOLOGY, AND SOCIETY. Several international monitoring projects are gathering data to help us understand coral reef destruction. In your opinion, why is it important to take action to protect coral reefs? Additional questions are available in CengageNOW at www.cengage.com/ login.

32

The Deuterostomes

© Ed Robinson/Tom Stack & Associates

Representative deuterostomes. This marine habitat photographed in Hawaii includes an echinoderm known as a pencil urchin (Heterocentrotrus mammilatus); and chordates, represented by fishes, the clown wrasse (Coris gaimard) with its yellow tail and the Moorish idol (Zanchus cornutus).

KEY CONCEPTS

W

hat does a sea star have in common with a fish, frog, hawk, or human? You may think it strange to group the echinoderms—the sea

32.1 Echinoderms and chordates are the two most successful deuterostome lineages in terms of diversity, number of species, and number of individuals.

stars, sea urchins, and sand dollars—with the chordates, the phylum to

32.2 Echinoderms are characterized by radial symmetry in

and morphological, developmental, and molecular data suggest that chor-

adults, a water vascular system, tube feet, and spiny skin.

32.3 At some time in its life, a chordate has a notochord;

which we humans and other animals with a backbone belong. However, even though these animals look and behave very differently, fossil evidence dates and echinoderms share a common ancestor and are closely related. In Chapter 31 we discussed the two major protostome branches of

a dorsal, tubular nerve cord; a muscular postanal tail; and an endostyle (or thyroid gland).

the animal kingdom, the lophotrochozoans and the ecdysozoans. In this

32.4 Invertebrate chordates include tunicates and lancelets.

chapter we focus on the third major branch of the animal kingdom, the

32.5 Shared derived characters of vertebrates include a

deuterostomes. Some biologists speculate that the last common ancestor of the deuterostomes was an animal that obtained food by filtering

vertebral column, cranium, neural crest cells, and an endoskeleton of cartilage or bone.

ocean water. The two major groups (phyla) of living animals assigned

32.6 Among the earliest vertebrates, ostracoderms were

to the deuterostomes are the echinoderms and chordates. The largest

armored, jawless fishes. The extant hagfishes and lampreys have neither jaws nor paired fins.

chordate subphylum is Vertebrata, which includes the animals with which

32.7 Jaws and fins were key adaptations that contributed to the success of jawed fishes, and the development of limbs was important in the transition to terrestrial life.

32.8 The evolution of the amniotic egg allowed animals to become fully terrestrial; the amniotic egg was the key adaptation leading to the evolution of reptiles, and later,

we are most familiar—fishes, amphibians, reptiles (including birds), and mammals. Echinoderms are represented in the photograph by the pencil urchin, and chordates are represented by the fishes. We begin this chapter with an introduction to the deuterostomes, and a survey of the echinoderms. With their unusual symmetry, spiny

skins, water vascular system, and tube feet, echinoderms are one of the most unique animal groups. We then turn our attention to the chordates. We describe the key chordate characteristics and discuss the invertebrate chordates. After an introduction to the vertebrates, we discuss jawless fishes and the evolution of jaws

32.2 ECHINODERMS ■ ■ LEARNING OBJECTIVE 2

Identify three shared derived characters of echinoderms and describe the main classes of echinoderms.

and limbs. We discuss evolution of the adaptations that made the transition to life on land possible. We then describe the evolution of reptiles (including birds) and mammals.

32.1 WHAT ARE DEUTEROSTOMES? ■ ■ LEARNING OBJECTIVE 1

Identify shared derived characters of deuterostomes and briefly describe the hemichordates.

Deuterostomia, the third major branch of the animal kingdom, includes the echinoderms—the sea stars, sea urchins, and sand dollars—and the chordates, the phylum to which we humans and other vertebrates (animals with a backbone) belong. Biologists also classify the hemichordates, a small group of wormlike marine animals, as deuterostomes. Hemichordates have a three-part body including a proboscis, collar, and trunk. These animals also have a characteristic ring of cilia surrounding the mouth. The most familiar of the hemichordates are the acorn worms, animals that live buried in mud or sand. Deuterostomes evolved from a common ancestor during the Proterozoic eon more than 550 million years ago (mya). The major groups were present during the Lower Cambrian. Deuterostomes are characterized by several shared derived characters (synapomorphies), evolutionary novelties absent in their most recent common ancestor (see Chapter 23). Their defining characters are similarities in their patterns of development. For example, deuterostomes are characterized by radial, rather than spiral, cleavage (see Chapter 30). Their cleavage is indeterminate, which means the fate of their cells is fixed later in development than is the case in protostomes. In deuterostomes, the mouth does not develop from the blastopore as in protostomes. The blastopore of deuterostomes becomes the anus (or is located near the future site of the anus), and the mouth develops from a second opening at the anterior end of the embryo—thus the name deuterostome, which is derived from the Greek words for “second mouth.” Another characteristic of deuterostomes is the presence of pharyngeal slits (openings in the pharynx) at some time in their life cycle. Hemichordates and chordates have pharyngeal slits. Although there is evidence that some early echinoderms had pharyngeal slits, the extant (living) echinoderms have lost this character. Basal deuterostomes (members of the earliest deuterostome group to evolve) have a type of larva with a loop-shaped band of cilia used for locomotion. Deuterostomes have a true coelom.

Review ■

What are three shared derived characters of deuterostomes?

The echinoderms (phylum Echinodermata) have one of the most highly derived body plans in the animal kingdom. Echinoderm larvae are bilaterally symmetrical, ciliated, and free-swimming. However, during development the body reorganizes, and the adult exhibits pentaradial symmetry, in which the body is arranged in five parts around a central axis. Biologists have hypothesized that early echinoderms were sessile and that radial symmetry evolved as an adaptation to that lifestyle. Their radial symmetry allows these animals to respond effectively in every direction of their surrounding environment. The most unique derived character of echinoderms is the water vascular system, a network of fluid-filled canals and chambers. In the sea star, for example, sea water enters through small pores in the madreporite, a sievelike structure on the body surface. Cilia lining the canals of the system move the water along. Branches of the water vascular system lead to numerous tiny tube feet that extend when filled with fluid. Each tube foot receives fluid from the main system of canals. A rounded muscular sac, or ampulla, at the base of the foot, stores fluid and is used to operate the tube foot. A valve separates each tube foot from other parts of the system. When the valve shuts, the ampulla contracts, forcing fluid into the tube foot. The fluid causes the tube foot to extend. At the bottom of the foot, a suction-type structure presses against and adheres to whatever surface the tube foot is on. The water vascular system functions in feeding and gas exchange, and serves as a hydrostatic skeleton important in locomotion. Another unique echinoderm character is the endoskeleton, an internal skeleton, covered by a thin, ciliated epidermis. The endoskeleton consists of calcium carbonate (CaCO3) plates and spines. The name Echinodermata, derived from words meaning “spiny skinned,” was inspired by the spines that project outward from the endoskeleton. Some groups have pincerlike, modified spines called pedicellariae on the body surface. These structures, found only among the echinoderms, keep the surface of the animal free of debris. Echinoderms have a well-developed coelom, containing coelomic fluid that transports materials. Although its structure varies in different groups, the complete digestive system is the most prominent body system. A variety of respiratory structures are found in the various classes. No excretory organs are present. The nervous system is simple, generally consisting of a nerve ring with nerves that extend out from it. Echinoderms have no brain. The sexes are usually separate, and eggs and sperm are generally released into the water, where fertilization takes place. The echinoderms evolved from bilaterally symmetrical ancestors, probably during the Early Cambrian period. They achieved maximum diversity by the middle of the Paleozoic era, about 400 mya. By the beginning of the Mesozoic era 251 mya, they had declined, leaving five main groups that have survived to the present day. Biologists have identified about 7000 living and more than

13,000 extinct species. All echinoderms inhabit marine environments. They are found in the ocean at all depths. We consider the five extant groups: class Crinoidea, sea lilies and feather stars; class

Asteroidea, sea stars; class Ophiuroidea, basket stars and brittle stars; class Echinoidea, sea urchins and sand dollars; and class Holothuroidea, sea cucumbers (FIG. 32-1 and TABLE 32-1).

Choanoflagellate ancestor

D. J. Wrobel/Monterey Bay Aquarium

Chordates

Hemichordates

Echinoderms

Ecdysozoans

Lophotrochozoans

Ctenophores

Cnidarians

Sponges

Deuterostomia

Marc Chamberlain/SeaPics.com

Robert Dunne/Photo Researchers, Inc.

(a) Class Crinoidea. Feather stars use their slender, jointed appendages to cling to the surface of a rock or coral reef. They can creep and often move away to escape predators.

(c) Class Ophiuroidea. Daisy brittle star (Ophiopholis aculeata), photographed in Muscongus Bay, Maine.

Peter Scoones/Seaphoto, Ltd.

D. J. Wrobel/Monterey Bay Aquarium

(b) Class Asteroidea. Orange and red sea star (Fromia monilis) on bubble coral.

(d) Class Echinoidea. With its flattened, circular body, the sand dollar (Dendraster excentricus) is adapted for burrowing on the ocean floor.

FIGURE 32-1 Representative echinoderms

(e) Class Holothuroidea. A sea cucumber (Thelonota) raises its body to spawn.

TABLE 32-1

Classes of Echinoderms

Class and Representative Animals

Characteristics

Crinoidea Feather stars, sea lilies

Feather stars motile; sea lilies, sessile, attach to ocean floor by stalk; oral surface on upper side of disc; tube feet along feathery arms trap microscopic organisms

Asteroidea Sea stars

Arms (rays) extend from central disc; tube feet on undersurface of each arm; mouth in center of underside of disc; carnivorous predators and scavengers

Ophiuroidea Basket stars, brittle stars

Resemble sea stars, but arms long, slender, and more set off from central disc; tube feet used to collect and handle food

Echinoidea Sea urchins, sand dollars

Skeletal plates flattened and fused to form a test; sand dollars burrow in sand and feed on organic particles; sea urchins graze on algae

Holothuroidea Sea cucumbers

Body is elongated, muscular sac; mouth surrounded by tentacles that are modified tube feet; sluggish animals that live on sea bottom; many species eject organs when conditions poor; for defense, can eject tubules out of anus

Feather stars and sea lilies are suspension feeders Feather stars and sea lilies (class Crinoidea) are the oldest living echinoderms (see Fig. 32-1a). Although many extinct crinoids are known, relatively few living species have been identified. The feather stars are motile, although they often remain in the same location for long periods. Sea lilies are sessile and remain attached to the ocean floor by a stalk. Crinoids remove suspended food from the water. Several branched, feathery arms extend upward. Numerous tube feet shaped like small tentacles are located along the feathery arms. These tube feet are coated with mucus that traps microscopic organisms. The oral surface (location of the mouth) is on the upper side of the disc. In all other echinoderms, the mouth is located on the underside of the disc toward the substratum.

Many sea stars capture prey Sea stars (commonly called starfish) are asteroids (class Asteroidea; see Fig. 32-1b). Their bodies consist of a central disc from which extend 5 to more than 20 arms, or rays (FIG. 32-2). The undersurface of each arm has hundreds of pairs of tube feet. The endoskeleton consists of a series of calcareous plates that permit some movement of the arms. Delicate dermal gills, small extensions of the body wall, carry on gas exchange. The mouth lies in the center of the underside of the disc.

Most sea stars are carnivorous predators and scavengers that feed on crustaceans, mollusks, annelids, and even other echinoderms. Occasionally, they catch small fish. The sea star’s water vascular system does not permit rapid movement, so its prey usually consists of stationary or slow-moving animals, such as clams. To attack a clam or other bivalve mollusk, the sea star mounts it and assumes a humped position as it straddles the edge opposite the hinge. Then, holding itself in position with its tube feet, the sea star slides its thin, flexible stomach out through its mouth and between the closed, or slightly gaping, valves (shell parts) of the clam. While the clam is still in its own shell, the sea star secretes enzymes that digest the soft parts of the clam to the consistency of a thick soup. When the partly digested meal is taken into the sea star body, it is further digested by enzymes secreted by digestive glands located in each arm. The circulatory system in sea stars is poorly developed and probably of little help in transporting materials. Instead, the coelomic fluid, which fills the large coelom and bathes the internal tissues, assumes this function. Metabolic wastes pass to the outside by diffusion across the tube feet and dermal gills. The nervous system consists of a ring of nervous tissue encircling the mouth and a nerve extending from this ring into each arm. Sea daisies are a curious group of asteroids characterized by small, disc-shaped, flat bodies (less than 1 cm in diameter) with no arms or mouth. They inhabit bacteria-rich wood sunk in deep water and apparently absorb bacteria through their body surface.

4

Organs have been removed to show digestive glands.

5

Cross section through arm Digestive gland

Madreporite Stomach Gonad Digestive gland

(b) Tube feet 1

Upper body and digestive glands have been removed, exposing ampullae of some of hundreds of tube feet (magnified view).

Upper surface with magnified detail. End is turned up to show tube feet on lower surface.

Spine

Gonad

Charles Seaborn/Odyssey Productions, Chicago

Anus

Ampulla

3

Tube feet

Tube feet

Dermal gill

2

(a)

Arm is dissected to show well-developed gonads.

Pedicellariae

(c)

5 mm

FIGURE 32-2 Body plan of a sea star (a) A sea star viewed from above, with its arms in various stages of dissection. Similar structures are present in each arm, but some organs are not shown in order to highlight certain structures. The two-part stomach is in the central disc with the anus on the aboral (upper) surface and the mouth beneath on the oral surface. (b) Cross section through arm and tube feet. (c) LM of tube feet of a sea star.

Basket stars and brittle stars make up the largest group of echinoderms Basket stars and brittle stars (serpent stars) make up the largest group of echinoderms (class Ophiuroidea) both in number of species and individuals (see Fig. 32-1c). These animals resemble sea stars in that their bodies consist of a central disc with arms. However, the arms are long and slender and more sharply set off from the central disc. Some brittle stars use their arms for locomotion, making rowing movements. Their tube feet lack suckers; they may be used to collect and handle food and may also serve a sensory function, perhaps that of taste.

Sea urchins and sand dollars have movable spines Sea urchins and sand dollars (class Echinoidea) have no arms (see Fig. 32-1d and chapter opening photograph). Their skeletal plates are flattened and fused to form a solid shell called a test. The flattened body of the sand dollar is adapted for burrowing in the sand, where it feeds on tiny organic particles. Sand dollars have smaller spines than sea urchins.

Many sea urchins graze on algae, scraping the sea floor with their calcareous teeth. Sea urchins use their tube feet for locomotion. They also push themselves along with their movable spines. In some species, the spines covering the body can penetrate flesh and are difficult to remove. So threatening are these spines that swimmers on tropical beaches are often cautioned to wear shoes when venturing offshore, where these living pincushions may dwell in abundance.

Sea cucumbers are elongated, sluggish animals Sea cucumbers (class Holothuroidea) are appropriately named, for some species are about the size and shape of a cucumber. The elongated body is a flexible, muscular sac (see Fig. 32-1e). The mouth is usually surrounded by a circle of tentacles that are modified tube feet. The endoskeleton consists of microscopic plates embedded in the body wall. The circulatory system, which is more highly developed than in other echinoderms, functions to transport oxygen and perhaps nutrients. Sea cucumbers are sluggish animals that usually live on the bottom of the sea, sometimes burrowing in the mud. Some graze with their tentacles, whereas others stretch their branched tentacles out

Vertebrates

Cephalochordates (lancelets)

Urochordates (tunicates)

Hemichordates (acorn worms)

Chordates

Echinoderms (sea stars, sea urchins)

in the water and wait for dinner to float by. Algae and other morsels are trapped in mucus along the tentacles. An interesting habit of some sea cucumbers is evisceration, in which the digestive tract, respiratory structures, and gonads are ejected from the body, usually when environmental conditions are unfavorable. When conditions improve, the lost parts are regenerated. Even more curious, when certain sea cucumbers are irritated or attacked, they direct their rear end toward the enemy and shoot red tubules out of their anus! These unusual weapons become sticky in sea water, and the attacking animal may become hopelessly entangled in the spaghetti-like tangle of tubules. Some of the tubules release a toxic substance.

Cranium, vertebral column, neural crest cells

Review ■ ■

What are three derived characters of echinoderms? Describe each. How do sea stars differ from crinoids? from sea urchins?

Notochord; dorsal, tubular nerve cord; postanal tail, endostyle

32.3 THE CHORDATES: DEFINING CHARACTERISTICS

Gill slits in pharynx Radial, determinate cleavage

■ ■ LEARNING OBJECTIVE 3

Deuterostome ancestor

Describe characteristics of chordates, including four shared derived characters.

FIGURE 32-3 Evolutionary relationships of the chordates

Biologists currently assign the chordates (phylum Chordata) to three groups, or subphyla: urochordates, marine animals called tunicates; cephalochordates, marine animals called lancelets; and vertebrates, animals with backbones. Systematists are currently debating whether the urochordates or cephalochordates are the sister group of the chordates (FIG. 32-3). Chordates are deuterostomes. They are coelomates with bilateral symmetry, a tube-within-a-tube body plan, and three welldeveloped germ layers. Four shared derived characters distinguish the chordates from all other groups of animals. These characters, which evolved in connection with their evolving methods of locomotion and obtaining food, include the notochord; the dorsal, tubular nerve cord; a postanal tail; and an endostyle or thyroid gland. 1. All chordates have a notochord during some time in their life cycle. The notochord is a dorsal, longitudinal rod composed of cartilage. The notochord is firm but flexible, and it supports the body. 2. At some time in their life cycle, chordates have a dorsal, tubular nerve cord. The chordate nerve cord differs from the nerve cord of most other animals in that it is located dorsally rather than ventrally, is hollow rather than solid, and is single rather than double. 3. Chordates have a larva or embryo with a muscular postanal tail, an appendage that extends posterior to the anus. 4. The endostyle is a groove in the floor of the pharynx that secretes mucus and traps food particles in the sea water passing through the pharynx. The endostyle is present in the urochordates, cephalochordates, and lamprey larvae. The thyroid gland evolved from the endostyle and is present in all other chordates.

This cladogram shows hypothesized phylogenetic relationships among deuterostomes, including chordates (tan region), based on structural and DNA data. The relative positions of the urochordates and cephalochordates may change as new data are considered.

Chordates have pharyngeal slits (also called pharyngeal gill slits) or pharyngeal pouches during some time in their life cycle (FIG. 32-4). In the embryo, a series of alternating branchial (gill) arches and grooves develop in the body wall in the pharyngeal (throat) region. Pharyngeal pouches extend laterally from the anterior portion of the digestive tract toward the grooves. In aquatic chordates, the tissue breaks through where the pouches meet the grooves, forming slits.

Brain

Dorsal, tubular nerve cord

Notochord

Mouth

Postanal tail

Pharynx Endostyle

Anus

Heart

Pharyngeal (gill) slits

Intestine

Muscular segments

FIGURE 32-4 Generalized chordate body plan Note the shared derived characters: notochord; dorsal, tubular nerve cord; postanal tail; and endostyle. Other key characteristics include pharyngeal (gill) slits and segmented muscles.

The perforated pharynx first evolved as an adaptation for filtering food. As water flowed through the mouth and passed out of the pharynx through the gill slits, suspended food particles were trapped in mucus. Later, modifications evolved, adapting the gill slits for gas exchange in fishes and some other aquatic vertebrates. Early in vertebrate evolution, anterior pharyngeal arches evolved into jaws. Pharyngeal slits are not a derived character of chordates because they are also present in hemichordates. Typically, chordates have an endoskeleton and a closed circulatory system with a ventral heart. Chordates have segmented bodies, but specialization is so pronounced that the basic segmentation of the body plan may not be apparent. For example, in vertebrates, although segmentation of the body is not obvious, muscles and nerves are segmentally organized.

Review ■ ■

What are four shared derived characters of chordates? How is the chordate nerve cord different from the nerve cord of most other animals?

32.4 INVERTEBRATE CHORDATES ■ ■ LEARNING OBJECTIVE 4 Compare tunicates and lancelets, and summarize the phylogeny of chordates.

Some chordates are not vertebrates. The invertebrate chordates include tunicates (urochordates) and lancelets (cephalochordates).

Tunicates are common marine animals The tunicates, or urochordates (subphylum Urochordata), include the sea squirts and their relatives (FIG. 32-5). Larval tunicates have typical chordate characteristics and superficially resemble tadpoles. The expanded body has a pharynx with slits, and the long muscular tail contains a notochord and a dorsal, tubular nerve cord. Some tunicates (appendicularians) retain their chordate features and ability to swim. These animals are common members of the zooplankton. Incurrent siphon Ganglion Oral tentacles

Excurrent siphon

Pharynx with slits

Tunic Vertebrates

Cephalochordates (lancelets)

Urochordates (tunicates)

Hemichordates (acorn worms)

Chordates

Echinoderms (sea stars, sea urchins)

Atrium Anus

Endostyle

Intestine

Esophagus

Testis

Digestive gland

Robert Shupak

Ovary

Deuterostome ancestor

Stomach

Heart

(a) Incurrent (top) and excurrent (side) siphons of a sea peach (Halocynthia aurantium), a solitary tunicate.

(b) Lateral view of an adult tunicate. The blue arrows represent the flow of water, and the red arrows represent the path of food.

Richard A. Cloney/ University of Washington

Pharynx with slits

Incurrent opening

Atrium Excurrent opening Nerve cord

Adhesive papilla

Notochord Heart

0.5 mm

(c) Swimming larval stage of a colonial species, Distaplia occidentalis.

FIGURE 32-5 Animated Tunicate body plan

Stomach

(d) Internal structure of a larval tunicate (lateral view).

Postanal tail

Lancelets clearly exhibit chordate characteristics Most cephalochordates (subphylum Cephalochordata) belong to the genus Branchiostoma, which consists of animals commonly known as lancelets, or amphioxus. Lancelets are translucent,

fish-shaped animals, 3 to 8 cm (1.3 to 3.2 in) long and pointed at both ends. They are widely distributed in shallow seas. Larvae can swim freely, but adults typically burrow in the sand in shallow water near the shore. In some parts of the world, lancelets are an important source of food. One Chinese fishery reports an annual catch of 35 tons (about one billion lancelets). Chordate characteristics are highly developed in lancelets. The notochord extends from the anterior tip (“head”; hence the name Cephalochordata) to the posterior tip. A dorsal, tubular nerve cord also extends the entire length of the animal, and many pairs of pharyngeal slits are evident in the large pharyngeal region (FIG. 32-6). Although superficially similar to fishes, lancelets have a far simpler body plan. They do not have paired fins, jaws, sense organs, a heart, or a well-defined head or brain. Like tunicates, lancelets are filter feeders. Cilia in the mouth and pharynx draw a current of water into the mouth. Microscopic organisms in the water are trapped in mucus secreted by the endostyle, and are then moved back to the intestine by beating cilia. Water passes through the pharyngeal slits into the atrium, a chamber with a ventral opening (the atriopore) located anterior to the anus. Metabolic wastes are excreted by segmentally arranged, ciliated protonephridia that open into the atrium. Unlike circulation in other invertebrates, the blood flows anteriorly in the ventral vessel and posteriorly in the dorsal vessel in lancelets. This circulatory pattern is similar to that of fishes.

Systematists debate chordate phylogeny Fossils of early chordates contribute to our understanding of chordate phylogeny. Invertebrate chordates are soft-bodied and did not leave many fossils. However, some well-preserved, key fossils that date back to the Middle Cambrian have been found in the Burgess Shale of British Columbia, Canada. Early Cambrian fossil sites have been discovered in Chengjiang and Haikou, China. At

Deuterostome ancestor

Vertebrates

Cephalochordates (lancelets)

Urochordates (tunicates)

Hemichordates (acorn worms)

Echinoderms (sea stars, sea urchins)

Chordates

Tentacles

(a)

Nerve cord

Tentacles

Notochord

Endostyle

Pharyngeal slits

Atrium

(b)

FIGURE 32-6 Animated Cephalochordate body plan (a) Photograph of a lancelet, Branchiostoma (amphioxus). Note the prominent pharyngeal gill slits. (b) Longitudinal section showing derived characters and other internal structures.

Intestine

Postanal tail Caudal fin

Gonads Atriopore

Anus

Dr. John Cunningham/Visuals Unlimited, Inc.

Most tunicates are ascidians, commonly known as sea squirts (class Ascidiacea). A sea squirt larva swims for a time, then attaches itself to a rock, piling, or to the sea bottom. It loses its tail, notochord, and much of its nervous system. Adult sea squirts are barrel-shaped, sessile marine animals unlike other chordates. Indeed, they are often mistaken for sponges or cnidarians. Only the pharyngeal slits, endostyle, and the structure of its larva indicate that the sea squirt is a chordate. Adult tunicates develop a protective covering, or tunic, that may be soft and transparent or quite leathery. Curiously, the tunic consists of a carbohydrate much like cellulose. The tunic has two openings: the incurrent siphon, through which water and food enter; and the excurrent siphon, through which water, waste products, and gametes pass to the outside. Sea squirts get their name from their practice of forcefully expelling a stream of water from the excurrent siphon when irritated. Tunicates are filter feeders that remove plankton suspended in the stream of water passing through the pharynx. Food particles are trapped in mucus secreted by cells of the endostyle, a groove that extends the length of the pharynx. Ciliated cells of the pharynx move the stream of food-laden mucus into the esophagus. Much of the water entering the pharynx passes out through the pharyngeal slits into an atrium (chamber) and is discharged through the excurrent siphon. Some species of tunicates form large colonies in which members share a common tunic and excurrent siphon. Colonial forms often reproduce asexually by budding. Sexual forms are usually hermaphroditic.

Chengjiang, fine-grained rocks about 530 million years old have preserved soft-bodied animals, including Haikouella, in some detail. Haikouella was about 2 to 3 cm (approximately 1 in) long and had a nerve cord, notochord, gills, muscle segments, and a brain. Many biologists view Haikouella as an early chordate. Some investigators hypothesize that this fossil represents a transitional form between cephalochordates and vertebrates. Pikaia, a lancelet-like fossil discovered in the Burgess Shale, had a primitive notochord with muscles attached to it that allowed for locomotion. About 4 cm (1.5 in) in length, Pikaia had a tail fin and probably filtered food from the water. Pikaia is considered an early chordate, most likely a cephalochordate. Are cephalochordates or urochordates the sister group of the vertebrates? Both structural and molecular data suggest that the cephalochordates are the sister group of the vertebrates. The most recent common ancestor of cephalochordates and vertebrates may have resembled a tunicate larva. Important shared derived characters include the presence of the notochord, nerve cord, and postanal tail in the adults. Another is the presence of somites, a series of paired blocks of mesoderm that develop on each side of the notochord. Somites are temporary structures that define the segmentation of the embryo. In vertebrates, they give rise to the vertebrae, ribs, and certain skeletal muscles. Another similarity between cephalochordates and vertebrates is the lancelet nerve cord, which has specialized regions, corresponding to the vertebrate forebrain and midbrain. Molecular data support the cephalochordates as the vertebrate sister group based on similarities in ribosomal and mitochondrial gene sequences. However, based on genome sequencing of tunicates (including the tunicate Ciona intestinalis), some biologists argue that the urochordate genome corresponds more closely to that of the chordate ancestor and that urochordates are the sister group of the vertebrates. The tunicate Ciona intestinalis is an excellent model organism for studying chordate development because its genome consists of only about 16,000 protein-coding genes, the basic set of genes found in chordates. Interestingly, about 80% of Ciona’s genes are found in vertebrates. In fact, Ciona has most vertebrate gene families, but in simplified form. For example, Ciona has only a single copy of each gene family involved in cell signaling and regulation of development, whereas vertebrates have two or more copies of these gene families. (A gene family is a group of genes that evolved over time from a single ancestral gene through duplication and divergence.) Some of the “extra” copies probably evolved to code for new structures or functions. Ciona also has genes for some vertebrate structures and processes, even though it does not express these genes. Although the debate about chordate phylogeny and the sister group of the vertebrates continues, in this edition of Biology, we consider the cephalochordates to be the sister group of the vertebrates.

Review ■

■

How are the main derived chordate characters evident in a tunicate larva and in an adult tunicate? If you found a small fishlike animal along the shoreline, how would you identify it as a lancelet?

32.5 INTRODUCING THE VERTEBRATES ■ ■ LEARNING OBJECTIVES 5 Describe four shared derived characters of vertebrates. 6 Describe the major taxa of extant vertebrates.

Numbering about 50,000 described species, the vertebrates (subphylum Vertebrata) are less diverse and much less numerous than the insects. However, vertebrates rival the insects in their adaptations to an enormous variety of lifestyles (FIG. 32-7). In addition to the basic chordate characteristics, vertebrates have a number of shared derived characters not found in other groups.

The vertebral column is a derived vertebrate character The vertebrates are distinguished from other chordates in having a backbone, or vertebral column, that forms the skeletal axis of the body. This flexible support develops around the notochord, and in most species it largely replaces the notochord during embryonic development. The vertebral column consists of cartilaginous or bony segments called vertebrae. Dorsal projections of the vertebrae enclose the nerve cord along its length. Anterior to the vertebral column, a cartilaginous or bony cranium, or braincase, encloses and protects the brain, the enlarged anterior end of the nerve cord. The cranium and vertebral column are part of the endoskeleton. In contrast with the nonliving exoskeleton of many invertebrates, the vertebrate endoskeleton is a living tissue that grows with the animal. Some vertebrates (jawless and cartilaginous fishes) have skeletons made of cartilage. However, in most vertebrates, the skeleton is mainly bone, a tissue that contains fibers made of the protein collagen. The hard matrix of bone consists of the compound hydroxyapatite, composed mainly of calcium phosphate. Many characters common to vertebrates have been derived from a group of cells called neural crest cells. These cells, found only in vertebrates, appear early in development and migrate to various parts of the embryo. Neural crest cells give rise to or influence the development of many structures, including nerves, head muscles, cranium, and jaws. Recall that Hox genes are important in determining pattern development in embryos (see Chapter 17). Specifically, Hox genes determine the fate of cells in the anterior-posterior axis. These genes occur in clusters on specific chromosomes. All invertebrates and early chordates that have been studied have one Hox gene cluster, but vertebrates have four duplicated clusters. Other gene clusters, including some encoding transcription factors, were also duplicated during the evolution of the early vertebrates. The significance of these genetic changes is the focus of current investigation, but it is reasonable to hypothesize that the increase in genetic complexity contributed to the development of more complex and varied bodies. Recall that invertebrates show an evolutionary trend toward cephalization, the concentration of nerve cells and sense organs

KEY POINT

This cladogram provides a framework for understanding the evolution of vertebrate diversity and adaptations to aquatic and terrestrial environments. Craniates V e

r

t e

b

r a

t e

s Tetrapods

Jawless fishes

Mammals

Reptiles (birds)

Reptiles

Amphibians

Dipnoi (lungfishes)

Actinistia (coelacanths)

Actinopterygii (ray-finned fishes)

Chondrichthyes (cartilaginous fishes)

Petromyzontida (lampreys)

Myxini (hagfishes)

Cephalochordates (lancelets)

Urochordates (tunicates)

Amniotes

Hair, mammary glands Limbs

Amniotic egg, adaptations that minimize water loss

Lungs (or swim bladder) Jaws Vertebrae Neural crest cells, head

Notochord, dorsal nerve cord, postanal tail, endostyle Chordate ancestor

FIGURE 32-7 Animated Evolutionary relationships of extant vertebrates This cladogram represents one phylogenetic interpretation of vertebrate phylogeny. The evolution of certain key characters is indicated. As new data are collected and considered, details of the cladogram will likely change. Figure 32-20 is a more detailed cladogram for amniote relationships.

in a definite head. Vertebrate evolution is characterized by pronounced cephalization. The brain has become larger and more elaborate, and its various regions have specialized to perform different functions. Either 10 or 12 pairs of cranial nerves emerge from the brain and extend to various organs of the body. Vertebrates have well-developed sense organs concentrated in the head: eyes; ears that serve as organs of balance and, in some vertebrates, for hearing as well; and organs of smell and taste.

Most vertebrates have two pairs of appendages. The fins of fishes are appendages that stabilize the fish in the water. Paired pectoral and pelvic fins are also used in steering. Biologists hypothesize that jointed appendages that facilitated locomotion on land evolved from the lobed (divided) fins of lungfishes. Vertebrates have a closed circulatory system with a ventral heart and blood containing hemoglobin. The complete digestive tract has specialized regions and large digestive glands (the liver

and pancreas). Several endocrine glands, which are ductless glands, secrete hormones. Paired kidneys regulate fluid balance. The sexes are typically separate. Early vertebrates were probably marine animals. In contrast to the tunicates and lancelets, which use cilia to beat a stream of water into the mouth and filter particles of food from the water, vertebrates use muscles for feeding. Some vertebrates use muscles to draw in a current of water from which both food and oxygen can be extracted. Muscles in the wall of the digestive tube are more powerful than cilia and probably contributed to an increase in both size and activity of early vertebrates. A muscular pharynx may have permitted animals to crush small prey after capture. The evolution of more effective sensory systems, a more complex brain, and organs that could support increased activity was important in the shift from filter-feeding to a more active lifestyle.

snakes, turtles, alligators and birds); and class Mammalia, the mammals. The reptiles and mammals form a clade known as amniotes, animals fully adapted to life on land.

Vertebrate taxonomy is a work in progress

Some of the earliest known vertebrates, collectively referred to as ostracoderms, consisted of several groups of small, armored, jawless fishes that lived on the bottom and strained their food from the water (see Fig. 21-11). Thick bony plates protected their heads from predators, and thick scales covered their trunks and tails. Most ostracoderms lacked fins. Fragments of ostracoderm scales have been found in rocks from the Cambrian period, but most ostracoderm fossils are from the Ordovician and Silurian periods. They became extinct by the end of the Devonian period. Like ostracoderms, present-day hagfishes and lampreys have neither jaws nor paired fins. They are eel-shaped animals, up to 1 m (about 3 ft) long. Their smooth skin lacks scales, and they are supported by a cartilaginous skeleton and well-developed notochord. Hagfishes, assigned to class Myxini, are marine scavengers (FIG. 32-8). They burrow for worms and other invertebrates or prey on dead and disabled fishes. Hagfishes use toothlike projections from their tongue to pull off flesh from prey. For leverage, a hagfish

■

■

What shared derived characters distinguish the vertebrates from the rest of the chordates? What are the six classes of fishes? the three classes of tetrapods?

32.6 JAWLESS FISHES ■ ■ LEARNING OBJECTIVE 7

Distinguish among the major groups of jawless fishes.

Tom McHugh/ Photo Researchers, Inc.

The study of evolutionary relationships of vertebrates is an important focus of research. The earliest vertebrate fossils include Haikouichthys, conodonts, and a group known as ostracoderms (discussed in the next section). Haikouichthys was a fishlike animal about 2.5 cm (1 in) long. It had several characters found in vertebrates, and some researchers think it was an early jawless fish. Conodonts were simple fishlike chordates with gill arches, muscular segments, and fins. They had large eyes, and complex toothlike hooks that were probably used in capturing prey. Conodonts were abundant from the Precambrian to the Late Triassic period; they have been identified in several fossil beds around the world. Cladistic analyses suggest that conodonts were early vertebrates. The earliest extant vertebrates to evolve were agnathans (a, “without”; and gnathos, “jaw”): the hagfishes, followed by the lampreys. Biologists assign hagfishes to class Myxini (mik-sin¿-y) and lampreys to class Petromyzontida (pet-tro-my-zon¿-tih-da) (from the Greek petros, “stone”; and myzon, “sucking”; lampreys hold onto stones by their mouths to prevent being washed away by the currents). Some systematists have argued that the hagfishes do not quite qualify as vertebrates. Although they have many vertebrate characters, including a cranium, hagfishes have no trace of vertebrae. The notochord is their only axial support. Some biologists use the term Craniata, based on the presence of a cranium, to designate a clade that includes the vertebrates plus the hagfishes. They view the Myxini as the sister group of all other craniates. Recent molecular analyses, however, support classifying hagfishes as vertebrates. Hagfishes may have evolved from vertebrates that had vertebrae but lost them during the course of their evolution. If we include the Myxini, the extant vertebrates are currently assigned to nine classes: six classes of fishes and three of tetrapods (four-limbed vertebrates) (TABLE 32-2 and see Fig. 32-7). Some biologists give fishes and tetrapods superclass status: superclass Pisces (fishes) and superclass Tetrapoda. In this classification scheme, the fishes include the hagfishes, lampreys, and four classes of jawed fishes: Chondrichthyes, Actinopterygii, Actinistia, and Dipnoi. The four-limbed land vertebrates, or tetrapods, include class Amphibia (frogs, toads, and salamanders); class Reptilia (lizards,

Review

FIGURE 32-8 Animated Pacific hagfish (Eptatretus stoutii) The hagfish has a flexible body and can tie it into knots. It knots its tail and then slides the knot toward its head. Being knotted gives it leverage when tearing flesh from its prey. Knotting itself also helps the hagfish remove excess slime from its body.

TABLE 32-2

Extant Vertebrate Classes

Class

Examples

Characteristics

Myxini

Hagfishes

Jawless, marine fishes that lack paired appendages; the notochord is their only axial support; lack vertebrae, so some biologists classify as craniates, but not as vertebrates

Petromyzontida

Lampreys

Jawless, freshwater and marine fishes with skeleton of cartilage; complete cranium and rudimentary vertebrae; hatch as small larvae

Chondrichthyes

Sharks, rays, skates, chimaeras

Jawed marine and freshwater fishes with skeleton of cartilage; notochord replaced by vertebrae in adult; gills; placoid scales; two pairs of fins; oviparous, ovoviparous, or viviparous (a few species); welldeveloped sense organs (including lateral line system)

Actinopterygii (ray-finned fishes)

Perch, salmon, tuna, trout

Bony, marine and freshwater fishes; gills; swim bladder; generally oviparous

Actinistia (lobe-finned fishes)

Coelacanths

Bony fishes; marine, nocturnal predators on fish; lobed fins

Dipnoi

Lungfishes

Bony freshwater fishes; four similarly sized limbs, which are similar in structure and position to those of tetrapods

Amphibia

Salamanders, frogs and toads, caecilians

Tetrapods; aquatic larva typically undergoes metamorphosis into terrestrial adult; gas exchange through lungs and/or moist skin; heart consists of two atria and single ventricle; systemic and pulmonary circulation

Reptilia

Turtles, lizards, snakes, alligators

Amniotes* with horny scales; adapted for reproduction on land (internal fertilization, leathery shell, amnion); lungs; ventricles of heart partly divided

Robins, pelicans, eagles, ducks, penguins, ostriches

Amniotes with feathers; anterior limbs modified as wings; compact, streamlined body; four-chambered heart; endothermic; vocal calls and complex songs

Monotremes (Protheria), marsupials (Metatheria), Eutheria (mammals with welldeveloped placentas)

Amniotes with hair; females nourish young with mammary glands; differentiation of teeth; three middle-ear bones; diaphragm; fourchambered heart; endothermic; highly developed nervous system

Mammalia

*A cladogram classification of amniotes is shown in Figure 32-20.

can tie itself into a knot. As a defense mechanism, hagfishes secrete large amounts of fluid that forms sticky slime in sea water. The hagfish becomes so slippery that predators cannot grasp it. Lampreys are jawless vertebrates assigned to class Petromyzontida. Some spend their adult lives in the ocean and return to fresh water to reproduce. Many species of adult lampreys are parasites on other fishes (FIG. 32-9). Adult parasitic lampreys have a circular sucking disc around the mouth, which lies on the ventral side

of the anterior end of the body. Using this disc to attach to a fish, the lamprey bores through the skin of its host with horny (made of keratin) teeth on the disc and tongue. Then the lamprey injects an anticoagulant into its host and sucks out blood and soft tissues. As in hagfish, the notochord persists throughout life and is not replaced by a vertebral column. However, lampreys have rudiments of vertebrae, cartilaginous segments called neural arches that extend dorsally around the spinal cord.

Courtesy of Dr. Kiyoko Uehara

Tom Stack/ Tom Stack & Associates

(a) Three lampreys attached to a carp by their suction-cup mouths. Note the absence of jaws and paired fins.

(b) Suction-cup mouth of adult lamprey (Entosphenus japonicus). Note the rasplike teeth.

FIGURE 32-9 Animated Lampreys Parasitic lampreys attach to fish and suck out body fluids. The wounds they leave in the fish may be fatal.

Review ■ ■

How do lampreys and hagfishes resemble ostracoderms? How do hagfishes differ from other fishes?

32.7 EVOLUTION OF JAWS AND LIMBS: JAWED FISHES AND AMPHIBIANS

(a) An acanthodian. Climatius was a spiny-skinned acanthodian with large fin spines and five pairs of accessory fins between the pectoral and pelvic pairs. Climatius was a small fish that reached a length of 8 cm (3 in).

■ ■ LEARNING OBJECTIVE 8

Trace the evolution of jawed fishes and early tetrapods; describe modern amphibians.

Imagine trying to hunt for food and then eat without jaws or limbs. This was the challenge for early vertebrates and helps explain why the jawless fishes are mainly scavengers and parasites. The evolution of jaws from a portion of the gill arch skeleton and the development of fins allowed fishes to become active predators. With jaws an animal can grasp and hold on to live prey while eating it. The evolution of fins allowed fishes to swim faster and with more control. Fishes with jaws and fins had many new opportunities for capturing food. Fossil evidence suggests that jaws and paired fins evolved during the Late Silurian and Devonian periods. Two early groups of jawed fishes, now extinct, were the acanthodians, armored fishes with paired spines and pectoral and pelvic fins, and placoderms, armored fishes with paired fins (FIG. 32-10). The success of the jawed vertebrates probably contributed to the extinction of the ostracoderms. When the fins of certain fishes evolved into limbs about 370 mya, a different type of locomotion was possible. These fishes

(b) A placoderm. Dinichthys was a giant placoderm that grew to a length of 8 m (26 ft). (Most placoderms were only about 20 cm, or 8 in, long.) Its head and thorax were covered by bony armor, but the rest of the body and tail were naked.

FIGURE 32-10 Early jawed fishes Acanthodians and placoderms flourished in the Devonian period.

were able to move about in shallow waters and wetlands in search of food. When the first tetrapods—the amphibians—moved onto the land, new opportunities for habitats and food became available.

Most cartilaginous fishes inhabit marine environments The cartilaginous fishes, members of class Chondrichthyes (kondrik¿-thee-eez), appeared as successful marine forms in the Devonian

Jeffrey L. Rotman / Peter Arnold, Inc.

period. Class Chondrichthyes , which is considered monophyletic, includes the sharks, rays, and skates (FIG. 32-11). Most species are ocean dwellers, but a few have invaded fresh water. With the exception of whales, the sharks are the largest living vertebrates. Some whale sharks (Rhincodon) exceed 15 m (49 ft) in length. Most rays and skates are flattened creatures that live partly buried in the sand. Their enormous pectoral fins propel them along the bottom, where they feed on mussels and clams. The stingray has a whiplike tail with a barbed spine at its base that can inflict a painful wound. The electric ray has electric organs on either side of the head. These modified muscles can discharge enough electric current (up to 200 volts) to stun fairly large fishes as well as human swimmers. The chondrichthyes retain their cartilaginous embryonic skeleton. Although this skeleton is not replaced by bone, a deposit of calcium salts may strengthen it. All chondrichthyes have paired jaws and two pairs of fins. The skin contains placoid scales, which are toothlike structures (FIG. 32-12). The lining of the mouth contains larger, but essentially similar, scales that serve as teeth. The teeth of other vertebrates are homologous with these scales. Shark teeth are embedded in the flesh and not attached to the jawbones; new teeth develop continuously in rows behind the functional teeth and migrate forward to replace any that are lost.

Kelvin Aitken /Peter Arnold, Inc.

(a) Blue-spotted stingray (Taeniura lymma ). Stingrays typically feed on shellfish and bottom-dwelling fishes.

(b) The great white shark (Carcharodon carcharias). Photographed in Australia, this shark is considered the most dangerous shark to humans. It is actually white only on its ventral aspect; the rest of the body is brownish gray or bluish gray.

FIGURE 32-11 Animated Cartilaginous fishes

The streamlined shark body is adapted for rapid swimming. Lift is provided by body shape and fins and by the action of swimming swiftly. The shark stores a great deal of oil in its large liver (which may account for up to 30% of the body weight). Fats and oils decrease the overall density of fishes and contribute to buoyancy. Even so, the shark body is denser than water, so sharks tend to sink unless they are actively swimming. Most sharks are predators that swim actively and catch and eat other fishes as well as crustaceans and mollusks. The largest sharks and rays, like the largest whales, are filter feeders that strain plankton from the water. They gulp water through the mouth. As the water passes through the pharynx and out the gill slits, food particles are trapped in a sievelike structure. Predatory sharks are attracted to blood, so a wounded swimmer or a skin diver towing speared fish is a target. However, despite the common portrayal of sharks in books and films as monstrous enemies, most do not go out of their way to attack humans. In fact, of the approximately 350 known shark species, fewer than 30 are known to attack humans. The shark has a complex brain and a spinal cord that is protected by vertebrae. The well-developed sense organs effectively locate prey in the water. Sharks may detect other animals electrically before sensing them by sight or smell. Electroreceptors on the shark’s head sense weak electric currents generated by the muscular activity of animals. The lateral line organ, found in all fishes, is a groove along each side of the body with many tiny openings to the outside. Sensory cells in the lateral line organ detect vibrations caused by waves and other movement in the water, including movements by predators or prey (see Fig. 43-7). Cartilaginous fishes have no lungs. Gas exchange takes place through their five to seven pairs of gills. A current of water enters the mouth and passes over the gills and out the gill slits, constantly providing the fish with a fresh supply of dissolved oxygen. Sharks that actively swim depend on their motion to enhance gas exchange. Rays, skates, and sharks that spend time on the ocean floor use muscles of the jaw and pharynx to pump water over their gills. The digestive tract of sharks consists of the mouth cavity; a long pharynx leading to the stomach; a short, straight intestine; and a cloaca, which opens on the underside of the body and is characteristic of many vertebrates (see Fig. 32-12). The liver and pancreas discharge digestive juices into the intestine. The cloaca receives digestive wastes as well as metabolic wastes from the urinary system. In females, the cloaca also serves as a reproductive organ. The sexes are separate, and fertilization is internal. In the mature male, each pelvic fin has a slender, grooved section, known as a clasper, used to transfer sperm into the female’s cloaca. The eggs are fertilized in the upper part of the female’s oviducts. Part of the oviduct is modified as a shell gland, which secretes a protective coat around the egg. Skates and some species of sharks are oviparous; that is, they lay eggs. Many species of sharks, however, are ovoviviparous, meaning their young are enclosed in eggs and incubated within the mother’s body. During development, the young depend on stored yolk for their nourishment rather than on transfer of materials from the mother. The young are born after hatching from the eggs. A few species of sharks are viviparous. Not only do the embryos develop within the uterus, but much of their nourishment is

Spleen

Kidney

Caudal fin

Stomach Testis

Gill slits

Clasper

Mouth

Pelvic fin

Cloaca

Intestine

Pancreas

Heart Liver

Pericardial cavity

Pharynx

(a) Internal structure of a shark.

Teeth

(b) Placoid scales. These tough scales protect the shark against predators. The base of each placoid scale is embedded in the skin. Placoid scales are modified teeth. They are covered with hard enamel.

(c) Teeth. Placoid scales are embedded in the flesh along the inner surface of the jaw cartilage where they function as teeth. Sharks have several rows of teeth. Old teeth are shed and quickly replaced.

FIGURE 32-12 Anatomy of a shark

delivered to them by the mother’s blood. Nutrients are transferred between the blood vessels in the lining of the uterus and the yolk sac surrounding each embryo.

The ray-finned fishes gave rise to modern bony fishes Although bony fishes appear earlier in the fossil record than cartilaginous fishes, both groups may have evolved about the same time, during the Devonian period. The two groups share many characteristics (such as continuous tooth replacement), but they also differ in important ways. Most bony fishes are characterized by a bony skeleton with many vertebrae. Bone has advantages over cartilage because it provides excellent support and effectively stores calcium. Most species have flexible median and paired fins, supported by long rays made of cartilage or bone. Overlapping, bony dermal scales cover the body. A lateral bony flap, the operculum, extends posteriorly from the head and protects the gills. Unlike most sharks, bony fishes are oviparous. They lay an impressive number of eggs and fertilize them externally. The ocean sunfish, for example, lays more than 300 million eggs! Of course,

most of the eggs and young become food for other animals. The probability of survival is increased by certain behavioral adaptations. For example, many species of fishes build nests for their eggs and protect them. During the Devonian period, the bony fishes diverged into two major groups: the Sarcopterygii and the ray-finned fishes, class Actinopterygii. Fossils of the earliest sarcopterygians date back to the Devonian period, about 400 mya. Lungs and fleshy, lobed fins characterized these fishes. The flexible fins were not supported by rays of bone except at their tips. Instead, the fins were supported by a group of bones with joints between them. These fins had muscles that could move them. Lobed fins may have evolved as an adaptation to life on the bottom of the sea. Fish could use the fins to push against the bottom. Early sarcopterygians evolved along two separate lines: the lungfishes (class Dipnoi) and lobe-finned fishes (class Actinistia). Both lineages have lobed fins. Three genera of lungfishes survive today in the rivers of tropical Africa, Australia, and South America. The ray-finned fishes underwent two important adaptive radiations. The first gave rise during the Late Paleozoic era to a group of fishes that are now mostly extinct. The second radiation began during the Early Mesozoic era and gave rise to the modern bony fishes.

Swim bladder

Nerve cord Dorsal fins

Kidney

Ureter

Caudal fin

Brain

Nostril Urinary bladder

Pharynx with gills Heart

Anal fin

Gonad Liver Stomach Pelvic fin

Intestine

Cloaca

FIGURE 32-13 Animated Perch, a representative bony fish The swim bladder is a hydrostatic organ that enables the fish to change the density of its body and remain stationary at a given depth. Pectoral fins (not shown) and pelvic fins are paired.

The common ancestor of the modern bony fishes had primitive lungs that could exchange gases in air. Lungs for gas exchange were retained by the lobe-finned fishes and lungfishes. In the ray-finned fishes, the lungs became modified as a swim bladder, an air sac that helps regulate buoyancy (FIG. 32-13). Bones and muscles are heavier than water, and without the swim bladder the fish would sink. By regulating gas exchange between its blood and swim bladder, a fish can control the amount of gas in the swim bladder and thus change the overall density of its body. This ability allows a bony fish, in contrast to a shark, to hover at a given depth of water without much muscular effort. There are more species of bony fishes than of any other vertebrate group. Biologists have identified about 49,000 living species of freshwater and saltwater bony fishes, of many shapes and colors (FIG. 32-14). Bony fishes range in size from that of Paedocypris progenetica, which has been measured at less than 8 mm (about 0.3 in) long, to that of the ocean sunfish (or Mola; see Fig. 20-16c), which may reach 4 m (13 ft) and weigh about 1500 kg (about 3300 lb). Paedocypris progenetica, native to Indonesia, is the smallest known vertebrate. The diversity of bony fishes may have resulted from the action of Hox genes. During the radiation of ray-finned fishes, entire chromosomes duplicated, resulting in one or more additional clusters of Hox genes. The zebrafish (see Fig.17-6d) has seven Hox clusters. These additional genes could have provided the genetic material for the evolution of the diverse species of ray-finned fishes.

Descendants of the lungfishes moved onto the land Biologists thought the lobe-finned fishes were extinct by the end of the Paleozoic era, so in 1938 the scientific community was very excited when a commercial fisherman caught one off the coast of South Africa. Since that time more than 200 specimens of these giant “living fossils” have been found in the deep waters off the southeastern coast of Africa, the Comores Islands, Madagascar,

and Indonesia (FIG. 32-15). These fishes, known as coelacanths, measure nearly 2 m (about 6 ft) in length. They are nocturnal predators on other fishes. Until the 1980s, the prevailing hypothesis was that coelacanths were the living fossil representatives of the group that gave rise to the land vertebrates. Then, a few investigators challenged this hypothesis with morphological data. The external nasal openings of fossil and living lungfishes suggested that they, rather than the coelacanths, are the ancestors of the land vertebrates. Other similarities between lungfishes and land vertebrates are tooth enamel and four limbs that have the same structure and position (FIG. 32-16). (Recall that the fins of lungfishes are also lobed.) During the 1990s, the lungfish hypothesis gained support from comparisons of mitochondrial DNA. More recently, underwater studies of coelacanths have shown that these fish use their muscular fins to slowly swim rather than to “walk” on the ocean bottom. In contrast, living lungfishes do use their fins to “walk” on the river bottom. Biologists now generally agree that ancestors of lungfishes, rather than coelacanths, gave rise to the tetrapods. Molecular data support the structural evidence that the limbs of tetrapods evolved from the lobed fins of fishes. Proteins encoded by certain regulatory genes, including Hox genes, have been found at the same developmental times and in the same locations in both limbs and fins. These findings indicate that limb and fin development are governed by the same genes. What drove the evolution of tetrapod limbs? One hypothesis suggests that during the frequent seasonal droughts of the Devonian period, swamps became stagnant or dried up completely. Fishes with lobed fins had a tremendous advantage for survival under those conditions. They were strikingly pre-adapted for moving onto the land. They had lungs for breathing air, and their sturdy, fleshy fins allowed them to “walk” along in shallow water. These fins could support the fish’s weight, so it could emerge onto dry land and make its way to another pond or stream. A more recent hypothesis holds that limbs evolved in a fully aquatic environment. Animals with more developed limbs could

David Hall / Photo Researchers, Inc.

Jeffrey L. Rotman / Peter Arnold, Inc.

(b) The parrotfish (Scarus gibbus). The parrotfish feeds on coral, grinds it in its digestive tract, and extracts the coralline algae. The fish eliminates a fine white sand. These fishes contribute to white sand beaches in many parts of the world. Parrotfish begin life as females and later become males.

move along in shallow water or creep through dense aquatic vegetation more efficiently than their lobe-finned ancestors. Tiktaalik was a transitional form between fishes and tetrapods (FIG. 32-17). Biologists consider Tiktaalik, which lived about 375 mya (during the Devonian period), a fish because it had scales and fins. However, Tiktaalik also had tetrapod features such as a movable neck and ribs that supported lungs. Acanthostega was one of the earliest known tetrapods. Fossils from the Devonian period show that this animal had four legs with well-formed digits. However, its limbs were not sufficiently strong for walking effectively on land. Acanthostega was an aquatic animal, not an amphibian. Ichthyostega is the earliest amphibian fossil found to date. This animal had strong legs supported by welldeveloped shoulder and hip bones and muscles. Natural selection favored animals such as Ichthyostega that could explore shallow wetlands and make their way onto dry land. The ability to move about on land, however awkwardly, provided

Norbert Wu/Minden Pictures/NGS Image Collection

(a) The porcupinefish (Diodon hystrix ). This fish swallows air or water to inflate its body, a strategy that discourages potential predators. Photographed in the Virgin Islands.

(c) Anglerfishes. In some clades of anglerfishes, the much smaller male attaches himself to a female with his sharp teeth and becomes a lifelong parasitic mate. Most species have at least one long spine projecting from the middle of the head. This “lure” is used as bait to attract prey. The more than 200 species of anglerfish are assigned to order Lophiliformes.

Mark Erdmann

Perennou Nuridsany/Photo Researchers, Inc.

FIGURE 32-14 Modern bony fishes

FIGURE 32-16 South American lungfish (Lepidosiren paradoxus) FIGURE 32-15 Diver swimming with coelacanth For many years, biologists thought that ancestors of this lobe-finned fish (Latimeria chalumnae) gave rise to the amphibians.

Lungfishes were common in Devonian and Carboniferous times (400 mya to 300 mya). South American lungfish have paired lungs on either side of the throat and can survive for long periods if the river they inhabit dries up.

Gerald and Buff Corsi / Tom Stack & Associates

Zina Deretsky/National Science Foundation

(a) Red dart frog (Dendrobates histrionicus). Poison dart frogs, native to Central and South America, are active during the day. Most are brightly colored. Many secrete toxins through their skin as a chemical defense against predators.

FIGURE 32-17 Animated Tiktaalik

access to new food sources. Terrestrial plants were already established, and terrestrial insects and arachnids were rapidly evolving. A vertebrate that could survive on land had less competition for food. However, success on land required the evolution of several major adaptations in addition to legs. As already discussed, the lungfishes had lungs for gas exchange. This was important because gills cannot function in air. Life on land also required the evolution of vertebrae and muscles strong enough to support the weight of the body in air. Body coverings and other mechanisms were needed to protect animals against the drying effect of air. Evolution of ears that could hear sound transmitted through air and olfactory mechanisms for detecting airborne odors contributed to the ability to find food and mates, and to avoid predators. At first, animals returned to the water to reproduce, but eventually adaptations evolved that allowed animals to reproduce in terrestrial environments. The early vertebrate experience on land was so successful that this lineage gave rise to the amphibians and, later, the reptiles, including birds, and the mammals.

Amphibians were the first successful land vertebrates Biologists classify modern amphibians (class Amphibia) in three orders. Order Urodela (“visible tail”) includes salamanders, mud puppies, and newts, all animals with long tails; order Anura (“no tail”) is made up of frogs and toads, with legs adapted for hopping; and order Apoda (“no feet”) contains the wormlike caecilians (FIG. 32-18). Although some adult amphibians are quite successful as land animals and live in dry environments, most return to the water to reproduce. Eggs and sperm are typically released in the water. Amphibians undergo metamorphosis, a transition from larva to adult. The embryos of most frogs and toads develop into larvae called tadpoles. These larvae have tails and gills, and most feed on aquatic plants. After a time, the tadpole undergoes metamorphosis, which is regulated by hormones secreted by the thyroid gland. During metamorphosis, gills and gill slits disappear, the tail

© Maximilian Weinzier/Alamy

This extinct fish, which grew to 2.75 m (9 ft), had limblike fins and other tetrapod features.

(b) Spotted salamander (Ambystoma maculatum). This salamander spends most of its time underground. In early spring spotted salamanders gather in woodland ponds to reproduce.

FIGURE 32-18 Modern amphibians

is resorbed, and limbs emerge. The digestive tract shortens, and food preference shifts from plant material to a carnivorous diet; the mouth widens; a tongue develops; the tympanic membrane (eardrum) and eyelids appear; and the eye lens changes shape. Many biochemical changes accompany the transformation from a completely aquatic life to an amphibious one. Several salamanders, such as the mud puppy Necturus, do not undergo complete metamorphosis; they retain many larval characteristics even when sexually mature adults. Recall from Chapter 20 that this is an example of paedomorphosis (see Fig. 20-17). This type of development permits these salamanders to remain aquatic rather than having to compete on land. The coloration of amphibians may conceal them in their habitat or may be very bright and striking. Many of the brightly colored species are poisonous (see Fig. 32-18a). Their distinctive colors warn predators that they are not encountering an ordinary amphibian. Some frogs camouflage themselves by changing color. Adult amphibians do not depend solely on their primitive lungs for the exchange of respiratory gases. Their moist, glandular skin, which lacks scales and is plentifully supplied with blood vessels, also serves as a respiratory surface. The numerous mucous glands within

the skin help keep the body surface moist, which is important in gas exchange. The mucus also makes the animal slippery, facilitating its escape from predators. Some amphibians have glands in their skin that secrete poisonous substances harmful to predators. The amphibian heart is divided into three chambers: two atria receive blood, and a single ventricle pumps it into the arteries. A double circuit of blood vessels keeps oxygen-rich and oxygenpoor blood partially separate. Blood passes through the systemic circulation to the various tissues and organs of the body. Then, after returning to the heart, it is directed through the pulmonary circulation to the lungs and skin, where it is recharged with oxygen. The oxygen-rich blood returns to the heart to be pumped out into the systemic circulation again. We discuss the comparative anatomy of the heart and circulation of various vertebrate classes in Chapter 44.

Review ■

■

From an evolutionary perspective, what is the significance of each of the following: (1) placoderms, (2) lungfishes, (3) ray-finned fishes, (4) Ichthyostega, and (5) Tiktaalik? What changes allow amphibians to move from aquatic life as a tadpole to terrestrial life as an adult?

32.8 AMNIOTES ■ ■ LEARNING OBJECTIVES 9 Describe three vertebrate adaptations to terrestrial life. 10 Describe the reptiles (including the birds) and give an argument for

Embryo

Amnion (protects embryo) Chorion (encloses embryo and other membranes) Allantois (stores wastes in reptiles and birds) Yolk sac (encloses yolk, provides nutrients)

Shell

Albumen (provides nutrients)

FIGURE 32-19 Animated An amniotic egg The amnion is a protective membrane surrounding the embryo. Other extraembryonic membranes are the yolk sac, allantois, and chorion.

eggs, their embryos have an amnion and other extraembryonic membranes. Another important adaptation to terrestrial life is a body covering that minimizes water loss. Such a covering presents another challenge because it severely decreases gas exchange across the body surface. This challenge has been met by the evolution of efficient lungs and circulatory systems for exchange of oxygen and carbon dioxide. Amniotes also have physiological mechanisms for conserving water. For example, much of the water filtered from the blood by the kidneys is reabsorbed in the kidney tubules to decrease fluid loss during excretion of metabolic wastes.

including the birds in the reptile clade.

11 Describe five key characters of mammals; contrast protherian (monotremes), metatherian (marsupials), and eutherian mammals, giving examples of animals that belong to each group.

The evolution of reptiles from ancestral amphibians required many adaptations that allowed them to become completely terrestrial. Evolution of the amniotic egg was an extremely important event because it allowed terrestrial vertebrates to complete their life cycles on land. The amniotic egg contains an amnion, a membrane that forms a fluid-filled sac around the embryo. The amnion provides the embryo with its own private “pond,” permitting independence from a watery external environment. In addition to keeping the embryo moist, the amniotic fluid serves as a shock absorber that cushions the developing embryo. The evolution of the amniotic egg is so important to the success of terrestrial vertebrates—reptiles (including birds) and mammals—that biologists refer to these animals as amniotes. Amniotes are a monophyletic group because they have a recent common ancestor that was itself an amniote. In addition to the amnion, the amniotic egg has three other extraembryonic (not part of the developing body itself) membranes: yolk sac, chorion, and allantois (FIG. 32-19). These membranes protect the developing embryo, store nutrients (yolk sac), carry on gas exchange (chorion and allantois), and store wastes (allantois). We discuss development of the extraembryonic membranes in Chapter 51. Although most extant mammals do not lay

Our understanding of amniote phylogeny is changing Biology instructors once taught that the three classes of amniotes were Reptilia, Aves (birds), and Mammalia. However, cladistic analysis determined that class Reptilia was not a monophyletic group without the birds. Because it included some, but not all, of its descendants, it was paraphyletic (see Fig. 23-7). For this reason, most biologists now classify the birds as a group of reptiles. Biologists hypothesize that the earliest amniotes resembled lizards. By the Late Carboniferous period, about 290 mya, amniotes had undergone an impressive adaptive radiation and diverged into two main branches: diapsids and synapsids (FIG. 32-20 and TABLE 32-3). The term diapsid refers to the two pairs of openings in the temporal bones that characterize the skulls of these animals. Synapsid skulls have one pair of temporal openings. The diapsids comprise all extant reptilian groups, including the birds and most extinct reptiles. The synapsids include the extinct therapsid reptiles and the mammals. A second great adaptive radiation of amniotes occurred during the Mesozoic era, which ended about 66 mya. During that time reptiles were the dominant terrestrial animals (see Chapter 21). In fact, the Mesozoic era is known as the “Age of Reptiles.” These animals had radiated into an impressive variety of ecological lifestyles (see Figs. 21-13, 21-14, and 21-15). Some could fly, others became marine, and many filled terrestrial habitats. Among the

KEY POINT

The diapsid reptiles gave rise to the birds, and the synapsids gave rise to the mammals. Note the five (including the birds) main branches of extant diapsids.

Mammals

Birds

Saurischian dinosaurs (extinct)

Pterosaurs (extinct)

Crocodilians e.g. crocodiles, alligators

Squamates e.g. snakes, lizards

Tuataras

Ichthyosaurs (extinct)

Turtles

Diapsids

ds si er ap Th

Hair, mammary glands, endothermy

sid s

si ap

Di ap

Diapsid skull

Sy n

Loss of diapsid skull openings

ds

”

ds

si

Ar ch

ap

os

n “A

au

rs

Endothermy

Skull with single pair of temporal openings Amniotic egg

Amniote ancestor

FIGURE 32-20 Evolutionary relationships of some amniotes This cladogram shows proposed evolutionary relationships among reptiles, birds, and mammals. Many of these relationships are currently being studied and may be reinterpreted.

major groups were the pterosaurs (ter¿-uh-sawr), the flying reptiles; the saurischian (saw-ris¿-kee-un) dinosaurs, a group that included Tyrannosaurus and Diplodocus; and the ornithischian (awr-nuh-this¿-kee-un) dinosaurs, including Triceratops. Some of the dinosaurs were among the largest land animals to ever walk on Earth. Some dinosaurs apparently traveled in social groups and took care of their young. Fossil evidence supports the hypothesis that at least some dinosaurs were endotherms, meaning they used metabolic energy to maintain a constant body temperature despite changes in the temperature of the environment. An advantage of endothermy is that it allows animals to be more active. The natural history and evolution of dinosaurs and other early reptiles are discussed in more detail in Chapter 21.

The reptiles were the dominant land animals for almost 200 million years. Then, toward the end of the Mesozoic era, many, including all the dinosaurs and pterosaurs, disappeared from the fossil record. In fact, more than half of all animal species became extinct at that time (see Chapter 21). As a result of this mass extinction, there are many more extinct reptiles than living species.

Reptiles have many terrestrial adaptations Many reptilian characters are adaptations to terrestrial life. The female reptile secretes a protective leathery shell around the egg, which helps prevent the developing embryo from drying out. The shell presents a challenge for reproduction because sperm can-

TABLE 32-3

Some Major Groups of Nonavian Reptiles

Subclass (Superorder/Order)

Important Characters

Subclass Diapsida

Skull with two pairs of temporal openings

Order Testudines (Chelonia) Turtles, terrapins, tortoises Superorder Lepidosauria Order Squamata Snakes, lizards, iguanas, amphisbaenians (worm lizards)

Enclosed in bony shell; beak of keratin instead of teeth; some can withdraw head and legs into shell; temporal openings lost (“anapsids”) Overlapping scales; most successful of modern reptiles Flexible armor of overlapping, horny scales, which are shed Lizards: Slender body, typically with four limbs; movable eyelids Snakes: Elongated body with no limbs; flexible, loosely jointed jaw; forked tongue protrusible

Order Sphenodonta Tuataras

Third eye under skin of forehead; only two extant species

Superorder Ichthyosauria (Extinct) Marine reptiles of Mesozoic

Body shaped like that of modern dolphins; large eyes; vertical tails

Superorder Archosauria

Mainly terrestrial; some specialized for flight; ventricle completely divided

Order Crocodilia Crocodiles, alligators, caimans, gavials

Four-chambered heart; closest extant relative to birds

Pterosauria (Extinct) Flying reptiles of Mesozoic

Membranous wings

Saurischia (Extinct) Mesozoic dinosaurs (Tyrannosaurus, Diplodocus) Birds descended from this lineage

Some were two-legged carnivores; others were four-legged herbivores

Ornithischia (Extinct) Mesozoic dinosaurs (Triceratops)

Bipedal and quadrupedal herbivores

Subclass Synapsida Therapsida (Extinct) Mammals descended from this lineage

not penetrate it. Fertilization must take place within the body of the female before the shell is added. In this process, the male uses a copulatory organ (penis) to transfer sperm into the female reproductive tract. An amnion surrounds the embryo as it develops within the protective shell. The hard, dry, horny scales that are part of the reptile’s skin retard drying—another adaptation to life on land. This scaly protective armor, which also protects the reptile from predators, is shed periodically. The dry reptilian skin does not allow effective gas exchange. Reptilian lungs are better developed than the saclike lungs of amphibians. Divided into many chambers, the reptilian lung provides an increased surface area for gas exchange. Most reptiles have a three-chambered heart that is more efficient than the amphibian heart. The single ventricle has a partition, though incomplete, that separates oxygen-rich and oxygen-poor blood, facilitating oxygenation of body tissues. In crocodiles the partition is complete, and the heart has four chambers. Like fishes and amphibians, extant reptiles generally lack metabolic mechanisms for regulating body temperature. They are

Skull with one pair of temporal openings Many mammal-like characters; became dominant land animals during Middle Permian

ectothermic, meaning that their body temperature fluctuates with the temperature of the surrounding environment. Some reptiles have behavioral adaptations that help them maintain a body temperature higher than that of their environment. For example, you may have observed a lizard basking in the sun, which raises its body temperature and so increases its metabolic rate. This permits the lizard to hunt actively for food. When the body of a reptile is cold, its metabolic rate is low and the animal tends to be sluggish. Ectothermy helps explain why reptiles are more successful in warm than in cold climates. Many reptiles are carnivorous (meat-eating). Their paired limbs, usually with five toes, are well adapted for running and climbing in search of prey. In addition, their well-developed sense organs enable them to locate prey.

We can assign extant reptiles to five groups Biologists assign the extant reptiles to five orders, or clades. Order Testudines includes the turtles, terrapins, and tortoises. Order

Turtles have protective shells

(a) Green turtle (Chelonia mydas). The paddlelike appendages of this turtle are adapted for swimming.

Frans Lanting / Minden Pictures

Members of order Testudines (te-stood¿-n-eez) are enclosed in a protective shell made of bony plates overlaid by horny scales. Some terrestrial species can withdraw their heads and legs completely into their shells. Their horny beak covers the jaws. Turtles do not have teeth.

Carlyn Iverson

Squamata includes lizards, snakes, and amphisbaenians (worm lizards). Order Sphenodonta includes the tuataras, lizardlike animals that live in burrows. The Archosaurs are a clade (sometimes designated a superorder) that includes the extinct flying reptiles and dinosaurs, as well as the extant order Crocodilia, and the birds. Order Crocodilia includes crocodiles, alligators, caimans, and gavials (FIG. 32-21).

Betty and Nathan Cohen /Visuals Unlimited

(b) The emerald tree boa (Corallus canina). The boa inhabits tropical South American forests. It rarely leaves the trees it inhabits to go to the ground. Emerald boas use their heat-sensitive pit organs to locate prey.

Smithsonian, Vol. 31, Feb. 2001/ Tom Blagden, Jr.

Pete Oxford / Minden Pictures

(d) Nile crocodile (Crocodilus niloticus). Both parents guard the leathery eggs. When the hatchlings emerge, the mother leads them to the water. She may carry them in her mouth.

(e) Two fledgling wood storks (Mycteria americana). About 8 to 9 weeks old, these storks preen their feathers atop their nest in South Carolina’s White Hall rookery.

(c) Tuatara (Sphenodon punctatus). These nocturnal predators reach a length of 60 cm (more than 2 ft) or longer. There are two species of tuataras. Both are classified as endangered.

FIGURE 32-21 Animated Representative extant reptiles

The size of adult turtles ranges in length from about 8 cm (3 in) to more than 2 m (6.5 ft) in leatherback turtles, which are the largest marine species. The weight of a leatherback turtle can exceed 500 kg (more than 1000 lb). The forelimbs of marine turtles are modified into flippers. Sea turtles migrate hundreds of miles from the beaches where they hatch to feeding grounds. Females return to the same beaches where they hatched to mate and nest. Most species of sea turtles are endangered as a result of human activities. Biologists sometimes refer to the aquatic forms as turtles and to the land species as tortoises. Freshwater types are sometimes called terrapins.

Lizards and snakes are the most common modern reptiles Lizards and snakes are assigned to order Squamata (squa-ma¿-tah). These animals have rows of scales that overlap like shingles on a roof, forming a continuous, flexible armor that is shed periodically. Lizards range in size from certain geckos, which weigh as little as 1 g (less than 0.1 oz), to the Komodo dragon of Indonesia, which may weigh 100 kg (220 lb). Their body sizes and shapes vary greatly. Some lizards are legless, for example, the “worm lizards” and the glass snake (which is really a lizard). Snakes are characterized by a flexible, loosely jointed jaw structure that lets them swallow animals larger than the diameter of their own jaws. These reptiles have elongated bodies with no legs, although pythons have vestigial hind-limb bones (see Fig. 18-17). Remember that although not all the tetrapods have four limbs, all evolved from four-limbed ancestors. Snake eyes are covered by a transparent scale. They do not have movable eyelids. Also absent are an external ear opening, a tympanic membrane (eardrum), and a middle-ear cavity. The snake uses its forked tongue as an accessory sensory organ for touch and smell. Chemicals from the ground or air adhere to the tongue. The snake rubs the tip of its tongue across the opening of a sense organ in the roof of the mouth that detects odors. Pit vipers and some boas also have a prominent pit organ on each side of the head that detects heat from endothermic prey (see Fig. 43-3). These snakes use their pit organs to locate and capture birds and small nocturnal mammals. Some snakes, such as king snakes, pythons, and boa constrictors, kill their prey by rapidly wrapping themselves around the animal and squeezing so it cannot breathe. Others have fangs, which are hollow teeth connected to venom glands. When the snake bites, it pumps venom through the fangs into the prey. Some snake venoms cause the breakdown of red blood cells; others, such as that of the coral snake, are neurotoxins, which interfere with nerve function. Venomous snakes of the United States include rattlesnakes, copperheads, cottonmouths, and coral snakes. All of these except the coral snakes are pit vipers. Amphisbaenians (am-fus-bee¿-nee-uns), or worm lizards, are now considered modified lizards. They are well adapted for their burrowing lifestyle. They have elongated, legless bodies, and their eyes are hidden under their skin. Most are less than 15 cm (6 in) long. Most species inhabit South America and southern Africa. Only one species (known as the “graveyard snake”) is known in the United States; it is found in Florida.

Tuataras superficially resemble lizards Tuataras look somewhat like iguanas but have certain distinct characters. For example, they are the only amniotes with vertebrae that are concave at both ends; such vertebrae are characteristic of fish and some amphibians. The two extant species of tuataras inhabit New Zealand and several small islands off the coast of New Zealand. Tuataras have long been considered endangered species. When humans brought rats, cats, and dogs to New Zealand, these non-native animals decimated the population of tuataras by eating their eggs. Tuataras became extinct on the New Zealand mainland. In 2005, these animals were reintroduced into a sanctuary in New Zealand and are now protected.

Crocodilians have an elongated skull The extant members of order Crocodilia (crok-uh-dil¿-ee-uh) along with the birds, are the surviving reptiles of the archosaur lineage. This is the lineage that gave rise to the pterosaurs (flying reptiles) and most of the dinosaurs that dominated the Mesozoic era. Modern crocodilians include three groups: (1) the crocodiles of Africa, Asia, and America; (2) the alligators of the southern United States and China, plus the caimans of Central America; and (3) the gavials of South Asia. Most species live in swamps, in rivers, or along seacoasts, feeding on various kinds of animals. Crocodiles are the largest living reptiles; some exceed 7 m (23 ft) in length. The cranial skeleton is adapted for aquatic life. The crocodile can be distinguished from the alligator or caiman by its long, slender snout and by the large fourth tooth on the bottom jaw that is visible when the mouth is closed.

How do we know that birds are really dinosaurs? Cladistic analyses indicate that birds evolved from the lineage of saurischian dinosaurs, specifically from the theropods (theer¿uh-pods), a group of bipedal, saurischian dinosaurs that included Tyrannosaurus and Deinonychus (see Fig. 21-15). This view is supported by the remarkable array of fossils discovered in the Yixian Formation in China. These fossils date back to the Early Cretaceous period (128 mya to 124 mya). Some dinosaurs had feathers Paleontologists have identified many theropod dinosaur fossils that demonstrate the evolution of feathers from very simple, tubular structures to complex modern forms. Caudipteryx, one feathered dinosaur discovered in the Yixian Formation, was about the size of a turkey. It had only a few teeth in the front of its upper jaw, and its bones show many birdlike characteristics (FIG. 32-22a). Caudipteryx has well-preserved imprints of complex feathers on its tail and forelimbs. Analyzing fossil evidence and molecular data, some investigators have hypothesized that feather evolution took place in terrestrial, bipedal dinosaurs before the evolution of birds or flight. Modern birds use asymmetrical feathers for flight. In 2003, paleontologists reported finding a new species of feathered dinosaur, Microraptor gui. Microraptor had asymmetrical feathers on both its

Early birds were transitional forms Many extinct theropods were long-tailed animals that moved about on two feet and had forelimbs with three clawed fingers. Although the bones of birds are fragile and disintegrate quickly, paleobiologists have found a few fossils of early birds. The first birds looked very much like dinosaurs. They had teeth (which modern birds lack), a long tail, and bones with thick walls. Certain extinct theropods and modern birds have feet with three digits and thinwalled, hollow bones. Both have a furcula, or wishbone, which is formed by the two clavicles (collarbones) fusing in the midline. Some biologists classify modern birds as theropods. The earliest known bird, Archaeopteryx (meaning “ancient wing”), was about the size of a pigeon. Ten specimens of this genus have been found in Bavaria in Jurassic limestone, which was laid down about 150 mya. Unlike those of extant birds, the jawbones of Archaeopteryx were armed with teeth, and its long reptilian tail was covered with feathers (FIG. 32-22b). Each of its short, broad wings had three claw-bearing, functional digits. Like modern birds, Archaeopteryx had wings, feathers, and a furcula. Its feathers were very similar to those of modern birds. Biologists do not know whether Archaeopteryx shared a common ancestor with other birds or whether Archaeopteryx itself gave rise to modern birds. Using computer tomography and computerized threedimensional reconstruction, researchers examined the inside of the Archaeopteryx skull. They found that Archaeopteryx had a birdlike brain with comparatively larger cerebral lobes than those of other reptiles. It had enlarged visual centers and highly developed inner ear canals similar to those of modern birds. These structures would support the coordination and agility needed for flight.

Pelvis

Leg

Chinese Academy of Geological Sciences

Feathers

(a) Caudipteryx exhibits both dinosaur and bird characteristics. This fossil lacks a head, but the rest of its skeleton is well preserved. The bones in the foot and the shape and orientation of the pelvis are similar to those of dinosaurs, but the fossil also has birdlike characters, including impressions of feathers.

From a painting by Rudolph Freund, courtesy of Carnegie Museum of Natural History

forelimbs and hind limbs, and also on its tail (See figure in Inquiring About: The Origin of Flight in Birds, in Chapter 21.) However, their small size and body structure support the hypothesis that flying evolved from gliding; that is, Microraptor climbed trees and used the feathered limbs for gliding. These small dinosaurs, which lived about 126 mya, coexisted with early birds. Biologists have debated why feathers may have evolved in dinosaurs. Were feathers important in courtship rituals? in camouflage? Were feathers an adaptation that conserved body heat? Insulating feathers could have contributed to the evolution of endothermy (the ability to maintain a constant body temperature), permitting animals to be more active. In 2010, paleontologists at George Washington University and their colleagues described a nearly complete skeleton of a basal theropod discovered in northwestern China’s Junggar Basin. This new fossil, named Haplocheirus sollers, was a feathered dinosaur that preceded the earliest known bird (Archaeopteryx) by about 15 million years. This finding supports the hypothesis that feather evolution took place in terrestrial, bipedal dinosaurs before the evolution of birds. Haplocheirus had organelles containing pigment that colored its feathers, suggesting that feathers first evolved for color displays in courtship rituals. The presence of feathers and the bone structure of Haplocheirus indicate that striking evolutionary convergence occurred between feathered dinosaurs and birds.

(b) This reconstruction represents the view that Archaeopteryx was a climbing animal that had at least some ability to use its wings and feathers for gliding.

FIGURE 32-22 Fossil of Caudipteryx, a feathered dinosaur, and reconstruction of Archaeopteryx, the earliest known bird

Gary Meszaros/Dembinsky Photo Associates

Alan G. Nelson/Dembinsky Photo Associates

© David Seawell/Alamy

(a) Southern cassowary (Casuarius casuarius), a ratite. The flightless southern cassowary, native to Australia, is the third largest bird in the world, after the ostrich and emu. This is the only bird known to have protective armor, including a hard crest that serves as a helmet atop its head.

(b) Male peacock (Pavo cristatus). The male peacock flares out its colorful feathers to get a female’s attention. Peafowl (peacocks and peahens) belong to the pheasant family.

(c) Atlantic puffin (Fratercula arctica). This puffin has caught several fish. The puffin’s beak becomes brightly colored during the breeding season.

FIGURE 32-23 Modern birds Although they are diverse, most birds are highly adapted for flight, and their basic structure is similar.

Cretaceous rocks have yielded fossils of other early birds. Hesperornis, which lived in North America, was a toothed, aquatic diving bird with small wings and broad, lobed feet for swimming. Ichthyornis was a toothed, flying bird about the size of a small tern. Based on fossils, structural similarities, and molecular data, most biologists now view birds as living dinosaurs.

Modern birds are adapted for flight By the end of the Cretaceous period, the major clades of presentday birds had evolved: palaeognaths and neognaths. (Some biologists assign birds to class Aves and consider palaeognathae and neognathae as superorders.) Palaeognaths (from Greek palaios, “ancient,” and gnathos, “jaw”) include flightless birds such as the ostriches, kiwis, cassowaries, and emus. These birds are called ratites. Their sternum (breastbone) is flat and does not have a ridge for attachment of wing muscles. They have only vestigial wings, but they have well-developed legs used for running. Most modern birds are neognaths (from Greek neos, “new,” and gnathos, “jaw”). These birds have powerful flight muscles attached to their keeled (ridged) sternum, and most neognaths fly. The penguins are an exception. They use their strong pectoral muscles and small, flipperlike wings to swim. Biologists have described about 9000 species of extant birds and classified them into about 23 monophyletic taxa. Birds are a diverse group (FIG. 32-23). They have become adapted to a variety of environments, and various species have different types of beaks,

feet, wings, tails, and behavioral patterns. They can be found on all continents, most islands, and even the open sea. The largest living birds are the ostriches of Africa, which may be up to 2 m (6.5 ft) tall and weigh 136 kg (300 lb). The great condors of the Americas have wingspans of up to 3 m (10 ft). The smallest known bird is the bee hummingbird of Cuba, with a length of less than 6 cm (about 2 in) and a weight of less than 2 g (about 0.06 oz). Like their reptilian ancestors, modern birds lay eggs and have reptilian-type scales on their legs. Birds have evolved remarkable specializations for flight. They are the only extant animals with feathers. Their feathers are an amazing example of biological engineering. They are very light, yet flexible and strong, and present a flat surface to the air. Feathers also protect the body and decrease water and heat loss. The anterior limbs of birds are wings, usually adapted for flight. The posterior limbs are modified for walking, swimming, or perching. Other adaptations for flight include the compact, streamlined body and the fusion of many bones, which provides the rigidity needed for flying. The bones are strong but very light. Many are hollow, containing large air spaces. The avian jaw is light, and instead of teeth, birds have a light, horny beak. The breastbone is broad and keeled for the attachment of the large flight muscles. Birds have lungs with air sacs, thin-walled extensions that occupy spaces between the internal organs and within certain bones. The unique “one-way” flow of air through their respiratory system is metabolically efficient because they do not need to pump air in and out of the lungs. (This respiratory system is described

in more detail in Chapter 46.) Birds have a four-chambered heart and a double circuit of blood flow. After blood delivers oxygen to the tissues, it is recharged with oxygen in the lungs; then the heart pumps the blood back into the systemic circulation. The very effective respiratory and circulatory systems provide the cells with enough oxygen to permit a high metabolic rate, which is necessary for the strenuous muscular activity that flying requires. Some of the heat a bird generates by metabolic activities helps it maintain a constant body temperature. Because they are endotherms, birds can remain active in cold climates. Birds excrete nitrogenous wastes mainly as semisolid uric acid. Because birds typically do not have a urinary bladder, these solid wastes are delivered into the cloaca. They leave the body with the feces, which are dropped frequently. This adaptive mechanism helps maintain a light body weight. Bills are specifically adapted for the type of food the bird eats (see Fig. 1-13). Birds must eat frequently because they have a high metabolic rate and typically do not store much fat. Although the choice of food varies widely among species, most birds eat energyrich foods such as seeds, fruits, worms, mollusks, or arthropods. Warblers and some other species eat mainly insects. Owls and hawks eat rodents, rabbits, and other small mammals. Some hawks catch snakes and lizards. Vultures feed on dead animals. Pelicans, gulls, terns, and kingfishers catch fish. An interesting feature of the bird digestive system is the crop, an expanded, saclike portion of the digestive tract below the esophagus, in which food is temporarily stored. The stomach is divided into a proventriculus, which secretes gastric juices; and a thick, muscular gizzard, which grinds food. The bird swallows small bits of gravel that act as “teeth” in the gizzard to mechanically break down food. Birds have a well-developed nervous system, with a brain that is proportionately larger than that of reptiles. Birds rely heavily on vision, and their eyes are relatively larger than those of other vertebrates. Hearing is also well developed. In striking contrast with the relatively silent reptiles, birds are very vocal. Most have short, simple calls that signal danger or influence feeding, flocking, or interaction between parent and young. Songs are usually more complex than calls and are performed mainly by males. Birds sing songs to attract and keep a mate and to claim and defend a territory. One of the most fascinating aspects of bird behavior is the annual migration made by many species. Some birds, such as the golden plover and Arctic tern, fly from Alaska to Patagonia, South America, and back each year, covering perhaps 40,250 km (25,000 mi) en route. Migration and navigation are discussed in Chapter 52. Many birds have beautiful, striking colors. The color is due partly to pigments deposited during the development of the feathers and partly to reflection and refraction of light of certain wavelengths. Many birds, especially females, are protectively colored by their plumage. During the breeding season, the male often assumes brighter colors to help him attract a mate.

Mammals have hair and mammary glands Hair is a key derived character of mammals (class Mammalia). All mammals have at least a few hairs at some time in their life, and no

other organism has true hair. Hair insulates the body, helping to maintain the high constant body temperature required for endothermy. The benefit of endothermy is a high metabolic rate, which allows a high level of activity even in low winter temperatures. Other derived characters of mammals are mammary glands, which produce milk for the young; one pair of temporal openings in the skull; differentiation of teeth into incisors, canines, premolars, and molars; and three middle-ear bones (malleus, incus, and stapes) that transmit vibrations from the tympanic membrane (eardrum) to the inner ear. The evolution of the cochlea, the organ of hearing in the inner ear, gives mammals an excellent sense of hearing. Contributing significantly to the success of the mammals, the nervous system is more highly developed than in any other group of animals. The cerebrum is especially large and complex, with an outer gray region, the cerebral cortex. In mammals, the highly specialized cerebral cortex, called the neocortex, has six layers of neurons. Specific regions in the neocortex are specialized for functions such as vision, hearing, touch, movement, emotional response, and higher cognitive functions. Fertilization in mammals is internal, and with the exception of the monotremes that lay eggs, mammals are viviparous. Most mammals develop a placenta, an organ of exchange between developing embryo and mother. As the mother’s blood passes through blood vessels in the placenta, it delivers nourishment and oxygen to the embryo and carries off wastes. By carrying their developing young internally, mammals avoid the hazards of having their eggs consumed by predators. By nourishing the young and caring for them, the parents offer both protection and an “education” on how to obtain food and avoid being eaten. A muscular diaphragm helps move air into and out of the lungs. Like birds, mammals are endotherms, but mammals may have evolved endothermy independently of birds. Some of the adaptations that allow mammals to maintain a constant body temperature include the covering of insulating hair, the four-chambered heart, and separation of pulmonary and systemic circulations. Red blood cells without nuclei serve as efficient oxygen transporters. The limbs of mammals are variously adapted for walking, running, climbing, swimming, burrowing, or flying. In four-legged mammals, the limbs are more directly under the body than they are in extant reptiles; this contributes to speed and agility.

New fossil discoveries are changing our understanding of the early evolution of mammals Mammals descended from therapsids, a group of synapsid reptiles, during the Triassic period more than 200 mya. The therapsids were doglike carnivores with differentiated teeth and legs adapted for running (FIG. 32-24). Until recently, early mammals were thought to be small, about the size of a mouse or shrew. However, fossil evidence indicates that mammals diversified earlier than previously thought. Some early mammals were fairly large, with limbs adapted for swimming or burrowing. Some had fur, and some were probably endothermic. How did mammals manage to coexist with reptiles during the approximately 160 million years that reptiles ruled Earth?

Painting by John C. Germann, Department of Library Services, American Museum of Natural History

small, early mammals had well-developed sense organs for hearing and smell. As many reptiles became extinct, mammals adapted to the niches (lifestyles) that reptiles abandoned. During this time, the flowering plants, including many trees, underwent an adaptive radiation, providing new habitats, sources of food, and protection from predators. Numerous new varieties of mammals evolved. During the Early Cenozoic era (more than 55 mya), the mammals underwent an adaptive radiation; they became widely distributed and adapted to an impressive variety of ecological lifestyles.

Modern mammals are assigned to three subclasses

FIGURE 32-24 Therapsid (Lycaenops) The therapsids were synapsid reptiles. Lycaenops, a carnivore about the size of a coyote, lived in South Africa during the Late Permian period.

Many adaptations permitted early mammals to compete for a place on this planet. Perhaps one of the most important was their skill at being inconspicuous. Many were arboreal (tree dwelling) and nocturnal (active at night), searching for food (mainly insects and plant material, and perhaps reptile eggs) at night while the reptiles were inactive. Although their eyes may have been

Marsupium

Eutheria

Metatheria (marsupials)

Protheria (monotremes)

Birds

Crocodilians (crocodiles, alligators)

Squamates (lizards, snakes)

Tuataras

Turtles

Mammalia

By the end of the Cretaceous period, three main groups of mammals had evolved (FIG. 32-25). Today, mammals inhabit virtually every corner of Earth; they live on land, in fresh and salt water, and in the air. They range in size from the tiny pygmy shrew, weighing about 2.5 g (less than 0.1 oz), to the blue whale, which may weigh up to 136,000 kg (150 tons) and which is probably one of the largest animals that has ever lived. Modern mammals are classified in two main clades: Protheria, which includes the egg-laying mammals, or monotremes, and Theria, which includes the mammals that bear their young alive. The therian clade is further divided into two groups: Metatheria, the marsupials (pouched mammals), and Eutheria, mammals that are more developed at birth than the marsupials. (The eutherians are often referred to as placental mammals because they have well-developed placentas. However, some marsupials do have simple placentas.) One group of protherians includes the duckbilled platypus and a second, the spiny anteaters or echidnas (FIG. 32-26; see also Figs. 1-16 and 23-5). These animals live in Australia and Tasmania; two species of spiny anteaters inhabit New Guinea. Female protherians lay eggs that they carry in a pouch on the abdomen or keep warm in a nest. When the young hatch, they lap up milk secreted by their mother’s mammary glands. Unlike other mammals, monotremes do not have nipples. With their long beaks and long, sticky tongues, spiny anteaters capture ants and termites. The duck-billed platypus,

ps na “A

Well-developed placenta Vivipary

s id p ia D ds si

si ap n Sy

Hair, mammary glands, endothermy

Tom McHugh/Photo Researchers, Inc.

”

ds si ap r e Th ds

Amniote ancestor

FIGURE 32-26 Spiny anteater FIGURE 32-25 Evolution of mammals This cladogram shows the current interpretation of the evolution of mammals.

The spiny anteater (Tachyglossus aculeatus) is an egglaying monotreme.

John Cancalosi/Peter Arnold, Inc. Robert Anderson, reprinted with permission of Hubbard Scientific Company

(a) Eastern gray kangaroo (Macropus giganteus) with young, known as a joey. The kangaroo is native to Australia.

(b) A kangaroo soon after birth. Marsupials are born in the embryonic state and continue to develop in the safety of the mother’s marsupium (pouch).

FIGURE 32-27 Marsupials

which lives in burrows along river banks, preys on freshwater invertebrates. It has webbed feet and a flat, beaverlike tail, which help it swim. Marsupials include pouched mammals such as kangaroos and opossums (FIG. 32-27a). Embryos begin their development in the mother’s uterus, where they are nourished by fluid and yolk. After a brief gestation, the young are born in an undeveloped stage. The young make their way to the marsupium (pouch), where they attach to a nipple. Nourished by the mother’s milk, they complete their development (FIG. 32-27b).

Eutherians

Marsupials

Lemur (Lemur )

Cuscus (Phalanger )

Anteater (Myrmecophaga )

Anteater (Myrmecabius )

Mouse (Mus )

Mouse (Dasycerus )

Flying squirrel (Glaucomys )

Flying phalanger (Phalanger )

Wolf (Canis )

Tasmanian wolf (Thylacinus )

Mole (Talpa )

Mole (Notoryctes )

Cat (Felis )

Cat (Dasyurus )

FIGURE 32-28 Convergent evolution in eutherian and marsupial mammals For each mammal in one group, there is a counterpart in the other group. Their similarities include both lifestyles and structural features.

At one time, marsupials probably inhabited much of the world, but placental mammals largely out-competed them. Now marsupials live mainly in Australia and in Central and South America. The only marsupial species that ranges into the United States is the Virginia opossum. Australia became geographically isolated from the rest of the world before placental mammals migrated there, and the marsupials remained the dominant mammals (see discussion of continental drift in Chapter 18). Australian marsupials underwent adaptive radiation, paralleling the evolution of placental mammals

James D. Watt/Animals Animals

McMurray Photography

(a) Wildebeest (Connochaetes taurinus), photographed in Tanzania. The dominant plains antelope in many areas of eastern and southern Africa, wildebeest migrate long distances during the dry season in search of food and water.

© Arco Images GmbH/Alamy

Michio Hoshino/Minden Pictures

(b) Humpback whales (Megaptera novaeangliae) exhibiting a rare double breach.

(c) Polar bears (Ursus maritimus), the largest living land carnivores, feed mainly on seals.

FIGURE 32-29 Eutherian mammals

elsewhere. Thus, marsupials that live in Australia and adjacent islands correspond to North American placental wolves, bears, rats, moles, flying squirrels, and even cats (FIG. 32-28 on facing page). Most familiar to us are the eutherian mammals, which are born at a more mature stage than marsupials. Indeed, among some species the young can walk and begin to interact with other members of the group within a few minutes after birth. Eutherians have a well-developed placenta (FIG. 32-29). The placenta develops from both embryonic membranes and the maternal uterine wall. In the placenta, blood vessels of the embryo come very close to the blood vessels of the mother, so materials can be exchanged by diffusion. (The two circulations do not normally mix.) The placenta allows the young to remain within the mother’s body until embryonic development is complete. Biologists assign extant placental

(d) Egyptian fruit bat (Rousettus aegyptiacus). Often called “flying foxes,” these small bats cluster in large colonies. They often hang by one foot from the cave ceiling.

mammals to about 19 orders. TABLE 32-4 gives a brief summary of some of these orders. Remember that aquatic mammals, such as dolphins, whales, and seals, evolved from terrestrial ancestors (see Fig. 18-9). Also refer to TABLE 32-5, which reviews the three main deuterostome groups.

Review ■ ■

■

■

What vertebrate adaptations are important for terrestrial life? Which amniotes are endothermic? What are some advantages of endothermy? What are some arguments for classifying birds and reptiles in a single clade? What are the three main clades of mammals? How do they differ?

TABLE 32-4

Some Orders of Extant Eutherian Mammals

Order, Representative Members, and Some Characteristics Insectivora (moles, hedgehogs, shrews) Nocturnal; eat insects. Shrew is smallest living mammal; some weigh less than 5 g.

African hedgehog Atelerix albiventris

Artiodactyla (cattle, sheep, pigs, deer, giraffes) Hoofed with even numbers of digits per foot; most have two toes, some have four. Most have antlers or horns. Herbivores; most are ruminants, chew a cud and have series of stomachs inhabited by bacteria that digest cellulose. American elk Cervus elaphus

Chiroptera (bats) Adapted for flying; fold of skin extends from elongated fingers to body and legs, forming wing. Guided in flight by type of biological sonar; emit high-frequency squeaks and are guided by echoes from obstructions. Eat insects and fruit or suck blood of other animals.

Carnivora (cats, dogs, wolves, foxes, bears, otters, mink, weasels, skunks, seals, sea lions, walruses) Carnivores with sharp, pointed canine teeth and molars for shearing; keen sense of smell; complex social interactions; among fastest, strongest, and smartest animals. Seals, sea lions, and walruses are large marine carnivores with limbs adapted as flippers for swimming.

Bat Eptesicus fuscus

Xenarthra (sloths, anteaters, armadillos) Teeth reduced or no teeth. Sloths are sluggish animals that hang upside down from branches; often protectively colored by green algae that grow on their hair. Armadillos are protected by bony plates; eat insects and small invertebrates.

Rodentia (squirrels, beavers, rats, mice, hamsters, porcupines, guinea pigs) Gnawing animals with chisel-like incisors. As they gnaw, teeth are worn down, and so must grow continually. Most numerous mammals both in numbers and species. Wolf Canis lupus

Nine-banded armadillo Dasypus bellus

Flying squirrel Glaucomys volans

Proboscidea (elephants) Largest land animals; weigh up to 7 tons; large head; broad ears; long, muscular, flexible trunk (proboscis); thick loose skin is characteristic; two upper incisors are elongated as tusks. Includes extinct mastodons and woolly mammoths.

Perissodactyla (horses, zebras, tapirs, rhinoceroses) Herbivores; hoofed with odd number of digits per foot; one or three toes; teeth adapted for chewing; usually large animals with long legs. Tapir Tapirus sp.

African elephant Loxodonta africana Lagomorpha (rabbits, hares, pikas) Like rodents, have chisel-like incisors; typically have long hind legs adapted for jumping; many have long ears. Pika Ochontona sp.

Sirenia (sea cows, manatees) Herbivorous, aquatic mammals with finlike forelimbs and no hind limbs. They are probably the basis for most tales about mermaids. Manatee Trichechus manatus

Primates (lemurs, monkeys, apes, humans) Highly developed brain and eyes; nails instead of claws; opposable thumb; eyes directed forward; omnivores; most species arboreal. (Primate evolution discussed in Chapter 22.)

Ring-tailed lemur Lemur catta

Cetacea (whales, dolphins, porpoises) Adapted for aquatic life with fish-shaped body and broad, paddlelike forelimbs (flippers); posterior limbs absent; many have thick layer of blubber under skin; some are filter feeders; mate and bear young in the water; very intelligent.

Humpback whale Megaptera noveangliae

TABLE 32-5

Overview of the Deuterostomes

Major Groups and Phyla

Body Plan

Key Characteristics

Echinoderms (sea stars, sea urchins; sand dollars)

Larva bilateral, ciliated; adult pentaradial; triploblastic; organ systems; complete digestive tube

Endoskeleton with spines; water vascular system functions in locomotion, feeding, and gas exchange; tube feet; marine

Hemichordates (acorn worms)

Bilateral symmetry; triploblastic; organ systems; complete digestive tube

Ring of cilia surrounds mouth; three-part body: proboscis, collar, and trunk; pharyngeal slits; wormlike marine animals

Chordates (tunicates, lancelets, vertebrates)

Bilateral symmetry; triploblastic; organ systems; complete digestive tube

Notochord; dorsal, tubular nerve cord; endostyle; pharyngeal slits during some time in life cycle; postanal tail; segmented body

Deuterostomes

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

32.1 (page 676)

are also characterized by pharyngeal (gill) slits, but this is a derived character of deuterostomes.

1 Identify shared derived characters of deuterostomes and briefly describe the hemichordates. ■ Deuterostomia includes echinoderms, hemichordates, and chordates. Shared derived characters include radial, indeterminate cleavage; the blastopore becomes (or is near the future site of) the anus; pharyngeal slits at some time in the life cycle. Basal deuterostomes have a larva with a loop-shaped ciliated band used for locomotion. ■ Hemichordates (acorn worms) are marine deuterostomes with a three-part body, including proboscis, collar, and trunk.

32.2 (page 676) 2 Identify three shared derived characters of echinoderms and describe the main classes of echinoderms. ■ Echinoderms (phylum Echinodermata) are marine animals with a spiny “skin,” water vascular system, tube feet, and endoskeleton. The larvae exhibit bilateral symmetry; most of the adults exhibit pentaradial symmetry. ■ Class Crinoidea includes sea lilies and feather stars. The oral surface of crinoids is turned upward; some crinoids are sessile. Class Asteroidea consists of the sea stars. They have a central disc ■ with five or more arms, and use tube feet for locomotion. ■ Class Ophiuroidea includes the brittle stars, which resemble sea stars but have longer, more slender arms that are set off more distinctly from the central disc. They use their arms for locomotion. Their tube feet lack suckers and are not used in locomotion. ■ Class Echinoidea includes the sea urchins and sand dollars. Echinoids lack arms; they have a solid shell and are covered with spines. Class Holothuroidea consists of sea cucumbers, animals with elongated flexible bodies. The mouth is surrounded by a circle of modified tube feet that serve as tentacles.

32.3 (page 680) 3 Describe characteristics of chordates, including four shared derived characters. ■ Phylum Chordata includes three subphyla: Urochordata, Cephalochordata, and Vertebrata. At some time in its life cycle, a chordate has a flexible, supporting notochord; a dorsal, tubular nerve cord; a muscular postanal tail; and an endostyle, or thyroid gland; they

32.4 (page 681) 4 Compare tunicates and lancelets, and summarize the phylogeny of chordates. ■ The tunicates, which belong to subphylum Urochordata, are suspension-feeding marine animals with tunics. Larvae have typical chordate characteristics and are free-swimming. Adults of most groups are sessile suspension feeders. ■ Subphylum Cephalochordata consists of the lancelets, small, segmented, fishlike animals; chordate characteristics are highly developed. ■ Biologists are still seeking evidence that will help determine whether the cephalochordates or the urochordates are the sister group of the vertebrates. Nerve cord

Notochord

Pharyngeal slits

Postanal tail

Endostyle

Learn more about the body plans of the deuterostomes by clicking on the figures in CengageNOW.

32.5 (page 683) 5 Describe four shared derived characters of vertebrates. ■

The vertebrates have a vertebral column composed of vertebrae that forms the chief skeletal axis of the body and a braincase, or cranium. Neural crest cells are important in the development of many structures, including the cranium and jaws. Vertebrates have pronounced cephalization, a complex brain, and muscles attached to the endoskeleton for movement. Explore the evolutionary relationships of vertebrates by clicking on the figure in CengageNOW.

Fertilization is internal. Amniotes have a body covering that retards water loss and physiological mechanisms that conserve water.

6 Describe the major taxa of extant vertebrates. ■

■

Vertebrates can be assigned to nine clades. The hagfishes, which make up the Myxini, and the lampreys, the Petromyzontida, have neither jaws nor paired fins. The Chondrichthyes comprise the sharks, rays, and skates; they are jawed fishes with skeletons of cartilage. The bony fishes are assigned to three groups: Actinopterygii, ray-finned fishes; Actinistia, coelacanths; and Dipnoi, lungfishes, which have limbs similar in size and position to the tetrapods. The tetrapods include the Amphibia (salamanders, frogs, and toads), which have aquatic larvae that undergo metamorphosis; Reptilia (turtles, lizards, snakes, alligators, and birds), amniotes with horny scales or feathers and reproduction adapted for terrestrial life; and Mammalia (monotremes, marsupials, placental mammals), amniotes with hair and mammary glands. (See Table 32-2 for a more detailed review.)

Learn more about the amniotic egg by clicking on the figure in CengageNOW.

32.6 (page 685) 7 Distinguish among the major groups of jawless fishes. ■

The extant jawless fishes, or agnathans, are the hagfishes (Myxini) and the lampreys (Petromyzontida). Jaws and paired fins are absent in both hagfishes and lampreys. Hagfishes are marine scavengers that secrete slime as a defense mechanism. Many lampreys are parasites on other fishes.

32.7 (page 687) 8 Trace the evolution of jawed fishes and early tetrapods; describe modern amphibians. Chondrichthyes, the cartilaginous fishes (sharks, rays, and skates), evolved during the Devonian period. They have jaws, two pairs of fins, and placoid scales. ■ During the Devonian period, bony fishes gave rise to two evolutionary lines: the Actinopterygii, or ray-finned fishes, and the Sarcopterygii, or lobe-finned fishes. The ray-finned fishes gave rise to the modern bony fishes. Their lungs have been modified as a swim bladder, an air sac for regulating buoyancy. ■ The Sarcopterygii probably gave rise to the lungfishes (Dipnoi) and the coelacanths (Actinistia). Evidence suggests that the lungfishes gave rise to the tetrapods, the land vertebrates. Tiktaalik was a transitional form between fishes and tetrapods. ■ Early amphibians (Amphibia) were mainly aquatic animals that ventured onto the land to find food or escape predators. These early tetrapods had limbs strong enough to support the weight of their bodies on land. Modern amphibians include salamanders, frogs and toads, and ■ caecilians. Most amphibians return to the water to reproduce. Frog embryos develop into tadpoles, which undergo metamorphosis to become adults. Amphibians use their moist skin as well as lungs for gas exchange. They have a three-chambered heart and systemic and pulmonary circulations. ■

Learn more about jawless, early jawed, cartilaginous, and bony fishes by clicking on the figures in CengageNOW.

32.8 (page 693) 9 Describe three vertebrate adaptations to terrestrial life. ■

Terrestrial vertebrates, or amniotes, include reptiles (including birds) and mammals. The evolution of the amniotic egg with its shell and amnion was an important adaptation for life on land. The amnion is a membrane that forms a fluid-filled sac around the embryo.

10 Describe the reptiles (including the birds) and give an argument for including the birds in the reptile clade. Biologists classify amniotes in two main groups: diapsids and synapsids. Diapsids include the turtles, ichthyosaurs, tuataras, squamates (snakes and lizards), crocodiles, pterosaurs, saurischian dinosaurs, and birds. The synapsids gave rise to the therapsids, which gave rise to the mammals. ■ Extant reptiles (including birds) are classified in five groups: turtles, terrapins, and tortoises; lizards, snakes, and amphisbaenians; tuataras; crocodiles, alligators, caimans, and gavials; and birds. ■ Reptiles have dry skin with horny scales, lungs with many chambers, and a three-chambered heart with some separation of oxygenrich and oxygen-poor blood. ■ Birds have many adaptations for powered flight, including feathers; wings; and light, hollow bones containing airspaces. They have a four-chambered heart and very efficient lungs, and they excrete solid metabolic wastes (uric acid). They are endotherms; that is, they maintain a constant body temperature. Birds have a welldeveloped nervous system with excellent vision and hearing. ■ Based on fossil evidence and molecular data, feather evolution took place in terrestrial, bipedal dinosaurs before the evolution of birds or flight. Birds are considered feathered reptiles that evolved from the lineage of saurischian dinosaurs, specifically from the theropods, a group of bipedal, saurischian dinosaurs. Like theropods, modern birds have feet with three digits; thin-walled, hollow bones; and a furcula, or wishbone. 11 Describe five key characters of mammals; contrast protherians (monotremes), metatherians (marsupials), and eutherian mammals, giving examples of animals that belong to each group. ■ Mammals have hair, mammary glands, differentiated teeth, and three middle-ear bones. They are endotherms and have a highly developed nervous system and a muscular diaphragm. ■ Protherians, the monotremes, include the duck-billed platypus and spiny anteaters. Monotremes lay eggs. ■ Metatherians, the marsupials, include pouched mammals, such as kangaroos and opossums. The young, born at an immature stage, complete their development in their mother’s marsupium, where they are nourished with milk from mammary glands. ■ Eutherians are more developed at birth than marsupials; they are characterized by a well-developed placenta, an organ of exchange that develops between the embryo and the mother. ■

T E S T YO U R U N D E R S TA N D I N G 1. Which of the following is not a shared derived character of echinoderms? (a) water vascular system (b) notochord (c) tube feet (d) pentaradial symmetry in adult (e) endoskeleton of calcium carbonate plates and spines 2. Which of the following organisms belongs to subphylum Vertebrata? (a) lophophorate (b) lamprey (c) lancelet (d) tunicate (e) crinoid 3. Which of the following is/are found in tunicates? (a) dorsal, tubular nerve cord (b) tube feet (c) anal gill slits (d) two pairs of appendages (e) vertebral column

10. Which of the following is a shared derived character of mammals? (a) placenta (b) amnion (c) three middle-ear bones (d) endostyle (e) notochord 11. Which of the following is not true? (a) feathers evolved in dinosaurs (b) birds are endothermic (c) birds excrete uric acid (d) birds have an amniotic egg (e) Archaeopteryx was the ancestor of feathered dinosaurs 12. Label the cladogram. See Figure 32-25 to check your answers.

Birds

4. Arrange the following animals to reflect an accepted sequence of evolution. 1. lungfish 2. dinosaur 3. amphibian 4. bird (a) 3, 1, 2, 4 (b) 1, 3, 2, 4 (c) 2, 1, 3, 4 (d) 1, 2, 3, 4 (e) 2, 4, 1, 3

Endothermy

5. A shark is characterized by (a) amnion (b) bony skeleton (c) water vascular system (d) placoid scales (e) placoid scales and amnion

Vivipary

6. Which of the following characteristics is associated with amphibians? (a) amnion (b) placoid scales (c) four-chambered heart (d) swim bladder (e) metamorphosis

s id

ps ra e Th

7. Reptiles (a) are endothermic (b) are amniotes (c) gave rise to the birds and mammals (d) have external fertilization (e) b and c are correct

si ap n Sy

8. Which of the following is not characteristic of birds? (a) hollow bones (b) amnion (c) ectothermy (d) high metabolic rate (e) reptilian-like scales on legs 9. Which of the following is not true of the duck-billed platypus? (a) classified as a marsupial (b) lays eggs (c) has a notochord at embryo stage (d) has vertebral column (e) is a predator

Hair, endothermy

ds

Amniote ancestor

CRITICAL THINKING 1. Sea urchins have radial symmetry. Explain why they are not classified as cnidarians.

5. EVOLUTION LINK. Most biologists now consider birds as living dinosaurs. Justify this position.

2. ANALYZING DATA. Imagine that you discover an interesting new animal. You find it has a dorsal, tubular nerve cord; a cranium; skin with scales; and a heart with two atria and two ventricles. How would you classify the animal? Explain each step in your decision. Where would you place this animal on the cladogram shown in Figure 32-20?

6. EVOLUTION LINK. Some biologists consider protherians (monotremes) to be therapsid reptiles rather than mammals. Give arguments for and against this position.

3. EVOLUTION LINK. Discuss the relationships among the echinoderms and chordates, describing shared derived characters that support grouping these animals as deuterostomes. 4. EVOLUTION LINK. Support the hypothesis that lungfishes gave rise to tetrapods.

7. SCIENCE, TECHNOLOGY, AND SOCIETY. Every year hundreds of sea turtles are injured and killed as a result of human activities. All seven species of marine turtles are listed under the Endangered Species Act. What measures would you propose to protect sea turtles? Additional questions are available in CengageNOW at www.cengage.com/ login.

33

Plant Structure, Growth, and Development

© Organica/Alamy

Foxglove, a herbaceous biennial. Purple and white foxgloves (Digitalis purpurea) are popular garden plants. Foxglove is also the source for digitalis, which is used to treat heart disease.

KEY CONCEPTS 33.1 The structure of plant cells, tissues, and organs matches their functions. The vascular plant body is differentiated into three tissue systems: the ground tissue system, the vascular tissue system, and the dermal tissue system. 33.2 Primary meristems lengthen roots and shoots throughout the lifespan of most plants. Woody plants have both primary meristems and secondary meristems, which increase roots and shoots in thickness.

T

he more than 300,000 species of flowering plants that live in and are adapted to the many environments on Earth represent remark-

able variety. Yet all of these—from desert cacti with enormously swollen stems, to cattails partly submerged in marshes, to orchids growing in the uppermost tree branches of lush tropical rain forests—are recognizable as plants. Almost all plants have the same basic body plan, which consists of roots, stems, and leaves. Plants are either herbaceous or woody. Herbaceous plants are nonwoody. In temperate climates, the aerial parts (stems and leaves) of her-

33.3 Plant development requires not only cell division and

baceous plants die back to the ground at the end of the growing season.

expansion but also cell determination and cell differentiation, and pattern formation and morphogenesis.

In contrast, the aerial parts of woody plants (trees and shrubs) persist. Botanically speaking, woody plants produce hard, lignified secondary tissues (the cell walls of secondary tissues contain lignin), and herbaceous

plants do not. Lignin and the production of secondary tissues are discussed later in this chapter and in Chapters 35 and 36. Annuals are herbaceous plants (such as corn, geranium, and marigold) that grow, reproduce, and die in one year or less. Other herbaceous plants (such as carrot, Queen Anne’s lace, cabbage, and foxglove) are biennials; they take two years to complete their life cycles before dying (see photograph). During their first season, biennials produce extra carbohydrates, which they store and use during their second year, when they typically form flowers and reproduce. Perennials are herbaceous and woody plants that have the potential to live for more than two years. In temperate climates,

The plant body of flowering plants (and other vascular plants) is usually organized into a root system and a shoot system (FIG. 33-1). The root system is generally underground. The aerial portion, the shoot system, usually consists of a vertical stem that bears leaves and, in flowering plants, flowers and fruits that contain seeds. Each plant typically grows in two different environments: the dark, moist soil and the illuminated, relatively dry air. Plants usually have both roots and shoots because they require resources from both environments. Thus, roots branch extensively through the soil, forming a network that anchors the plant firmly in place and absorbs water and dissolved minerals (inorganic nutrients). Leaves, the flattened organs of photosynthesis, are attached more or less regularly on the stem, where they absorb the sun’s light and atmospheric carbon dioxide (CO2) used in photosynthesis.

the aerial (aboveground) stems of herbaceous perennials such as iris, rhubarb, onion, and asparagus die back each winter. Their

The plant body consists of cells and tissues

underground parts (roots and underground stems) become dor-

As in other organisms, the basic structural and functional unit of plants is the cell. Plants have evolved a variety of cell types, each specialized for particular functions. Plant cells are organized into tissues. A tissue is a group of cells that forms a structural and functional unit. Simple tissues are composed of only one kind of cell, whereas complex tissues have two or more kinds of cells. In vascular plants, tissues are organized into three tissue systems, each of which extends throughout the plant body (FIG. 33-2). Each tissue system contains two or more kinds of tissues (TABLE 33-1). Most of the plant body is composed of the ground tissue system, which has a variety of functions, including photosynthesis, storage, and support. The vascular tissue system, an intricate conducting system that extends throughout the plant body, is responsible for conduction of various substances, including water, dissolved minerals, and food (dissolved sugar). The vascular tissue system also functions in strengthening and supporting the plant. The dermal tissue system provides a covering for the plant body. Roots, stems, leaves, flower parts, and fruits are organs because each is composed of all three tissue systems. The tissue systems of different plant organs form an interconnected network throughout the plant. For example, the vascular tissue system of a leaf is continuous with the vascular tissue system of the stem to which it is attached, and the vascular tissue system of the stem is continuous with the vascular tissue system of the root.

mant during the winter and send out new growth each spring. (In dormancy, an organism reduces its metabolic state to a minimum level to survive unfavorable conditions.) Similarly, in certain tropical climates with pronounced wet and dry seasons, the aerial parts of herbaceous perennials die back, and the underground parts become dormant during the unfavorable dry season. Other tropical plants, such as orchids, are herbaceous perennials that grow year-round. All woody plants are perennials, and some of them live for hundreds or even thousands of years. In temperate climates, the aerial stems of woody plants become dormant during the winter. Many temperate woody perennials are deciduous; that is, they shed their leaves before winter and produce new stems with new leaves the following spring. Other woody perennials are evergreen and shed their leaves over a long period, so some leaves are always present. Because they have permanent woody stems that are the starting points for new growth the following season, many trees attain massive sizes. In this chapter we focus on the structure, growth, and development of flowering plants, which are vascular plants characterized by flowers, double fertilization, endosperm, and seeds enclosed within fruits (see Chapter 28). We then examine other aspects of flowering plants in Chapters 34 to 38.

33.1 THE PLANT BODY ■ ■ LEARNING OBJECTIVES 1

2 3 4

Discuss the functions of various parts of the vascular plant body, including the nutrient- and water-absorbing root system and the photosynthesizing shoot system. Describe the structure and functions of the ground tissue system (parenchyma tissue, collenchyma tissue, and sclerenchyma tissue). Describe the structure and functions of the vascular tissue system (xylem and phloem). Describe the structure and functions of the dermal tissue system (epidermis and periderm).

The ground tissue system is composed of three simple tissues The bulk of a herbaceous plant is its ground tissue system, which is composed of three tissues: parenchyma, collenchyma, and sclerenchyma (TABLE 33-2 on pages 712 and 713). These tissues can be distinguished by their cell wall structures. Recall that plant cells are surrounded by a cell wall that provides structural support (see Chapter 4). A growing plant cell secretes a thin primary cell wall, which stretches and expands as the cell increases in size. After the cell stops growing, it sometimes secretes a thick, strong secondary cell wall, which is deposited inside the primary cell wall— that is, between the primary cell wall and the plasma membrane (see Fig. 4-31). Plant cell walls have several important roles. As discussed later in the chapter, cell walls are involved in growth; the ability of

KEY POINT

During the evolution of plants, the root and shoot systems specialized to obtain resources from the soil and air, respectively.

Developing fruit

Nodes (areas of leaf and axillary bud attachment)

Shoot system

Axillary bud Internode (area between adjacent nodes)

Petiole Blade Leaf

Anther of stamen

Stem

Stigma of pistil Rosette of basal leaves

Ovary of pistil (consisting of two carpels)

Sepal Root system

Petal

Taproot

Branch roots

Flower

FIGURE 33-1 Animated The plant body A plant body consists of a root system, usually underground, and a shoot system, usually aerial. Shown is mouse-ear cress (Arabidopsis thaliana), a small plant in the mustard family that is a model plant for

primary cell walls to expand allows cells to increase in size. Scientists are increasingly aware that signal transduction takes place in plant cell walls because many carbohydrate and protein molecules in plant cell walls communicate with other molecules both inside and outside the cell. The cell wall is also the first line of defense against disease-causing organisms.

biological research. Arabidopsis is native to North Africa and Eurasia and has been naturalized (was introduced and is now growing in the wild) in California and the eastern half of the United States.

Parenchyma cells have thin primary walls Parenchyma tissue, a simple tissue composed of parenchyma cells, is found throughout the plant body and is the most common type of cell and tissue (FIG. 33-3a on page 714). The soft parts of a plant, such as the edible part of an apple or a potato, consist largely of parenchyma.

KEY POINT

The tissue systems are continuous throughout the plant. For example, the vascular tissue system in a leaf is continuous with the vascular tissue system in the stem to which it is attached.

Dermal tissue system Vascular tissue system Ground tissue system

(a) Leaf Dermal tissue system Vascular tissue system Ground tissue system

(b) Stem Dermal tissue system Vascular tissue system Ground tissue system

(c) Root

FIGURE 33-2 Animated The three tissue systems in the plant body This figure shows the distribution of the ground tissue system, vascular tissue system, and dermal tissue system in a herbaceous eudicot such as Arabidopsis.

TABLE 33-1

Tissue Systems, Tissues, and Cell Types of Flowering Plants

Tissue System

Tissue

Cell Types

Main Functions of Tissue

Ground tissue system

Parenchyma tissue Collenchyma tissue Sclerenchyma tissue

Parenchyma cells Collenchyma cells Sclerenchyma cells (sclereids or fibers)

Storage, secretion, photosynthesis Support Support, strength

Vascular tissue system

Xylem

Tracheids Vessel elements Xylem parenchyma cells Fibers (sclerenchyma cells)

Conduction of water and nutrient minerals, support Conduction of water and nutrient minerals, support Storage Support, strength

Phloem

Sieve tube elements Companion cells Phloem parenchyma cells Fibers (sclerenchyma cells)

Conduction of sugar in solution, support May control functioning of sieve tube elements, loading sugar into sieve tube elements Storage Support, strength

Epidermis

Epidermal cells Guard cells Trichomes

Protective covering over surface of plant body Regulate stomata Variable functions

Periderm

Cork cells Cork cambium cells Cork parenchyma cells

Protective covering over surface of plant body Meristematic (form new cells) Storage

Dermal tissue system

TABLE 33-2

Cell Types in the Ground Tissue System

Phil Gates/Biological Photo Service

Nucleus

Parenchyma cell Description Living, actively metabolizing; thin primary cell walls Functions Storage; secretion, and/or photosynthesis Location and comments Throughout the plant body; shown is an LM of a stamen hair of a spiderwort (Tradescantia virginiana) flower; note the large pigmented vacuole; the nucleus is not inside the vacuole but is lying on top of the vacuole

Pigmented vacuole

Cell wall

Chloroplasts

Cell wall

Cengage

Vacuole

Parenchyma cell Description Living, actively metabolizing; thin primary cell walls Functions Storage; secretion; photosynthesis Location and comments Throughout the plant body; shown is an LM of leaf cells from an aquatic plant (Elodea); note the many chloroplasts in the thin layer of cytoplasm surrounding the large, transparent vacuole

Intercellular spaces

Cell wall

Cengage

Starch grains

Parenchyma cell Description Living, actively metabolizing; thin primary cell walls Functions Storage; secretion; photosynthesis Location and comments Throughout the plant body; shown is an LM of a cross section of part of a buttercup (Ranunculus) root; note the starch grains filling the cells

Ed Reschke/Peter Arnold, Inc.

Primary cell walls are thickened in corners

Collenchyma cell Description Living; unevenly thickened primary cell walls Function Elastic support Location and comments Just under stem epidermis; shown is an LM of a cross section of an elderberry (Sambucus) stem; note the unevenly thickened cell walls that are especially thick in the corners, making the cell contents assume a rounded shape in cross section

(continued)

TABLE 33-2

Nucleus

Cell Types in the Ground Tissue System (continued) Secondary cell walls

Cytoplasm

Pits

James Mauseth, University of Texas

Sclereid (sclerenchyma cell) Description May be living or dead at maturity; thick secondary cell walls; lacks secondary wall at pits Functions Strength; sclereid-rich tissue is hard and inflexible Location and comments Shells of walnuts and coconuts; pits of cherries and peaches; shown is an LM of living sclereids in a section through a cherry (Prunus avium) pit

James Mauseth, University of Texas

Thick secondary cell walls

Lumen where protoplast was when these cells were alive

Parenchyma cells perform several important functions, such as photosynthesis, storage, and secretion. Parenchyma cells that function in photosynthesis contain green chloroplasts, whereas nonphotosynthetic parenchyma cells lack chloroplasts and are often colorless. Materials stored in parenchyma cells include starch grains, oil droplets, water, and salts, which are sometimes visible as crystals. Resins, tannins, hormones, enzymes, and sugary nectar are examples of substances that may be secreted by parenchyma cells. The various functions of parenchyma require that parenchyma cells be alive and metabolically active. Parenchyma cells have the ability to differentiate into other kinds of cells, particularly when a plant has been injured. If xylem (water-conducting) cells are severed, for example, adjacent parenchyma cells may divide and differentiate into new xylem cells within a few days. (Recall from Chapter 17 that it is possible to induce certain plant cells to become the equivalent of embryonic stem cells that can then differentiate into specialized cells.) Collenchyma cells have unevenly thickened primary walls Collenchyma tissue, a simple plant tissue composed of collenchyma cells, is an extremely flexible structural tissue that provides much of the support in soft, nonwoody plant organs (FIG. 33-3b).

Fiber (sclerenchyma cell) Description Often dead at maturity; thick secondary cell walls; fewer pits than in sclereids Functions Support; provides strength Location and comments Throughout the plant body; common in stems and certain leaves; shown is an LM of a cross section through a clump of fibers from a century plant (Agave) leaf

Support is crucial for plants, in part because it lets them grow upward to compete with other plants for available sunlight in a plantcrowded area. Plants lack the bony skeletal system typical of many animals; instead, individual cells, including collenchyma cells, support the plant body. Collenchyma cells, which are usually elongated, are alive at maturity. Their primary cell walls are unevenly thickened and are especially thick in the corners. Collenchyma is not found uniformly throughout the plant and often occurs as long strands near stem surfaces and along leaf veins. The “strings” in a celery stalk, for example, consist of collenchyma tissue. Sclerenchyma cells have both primary walls and thick secondary walls Another simple plant tissue specialized for structural support is sclerenchyma tissue, whose cells have both primary and secondary cell walls. The root of the word sclerenchyma is derived from the Greek word sclero, meaning “hard.” The secondary cell walls of sclerenchyma cells become strong and hard because of extreme thickening. Pits, which are thin areas where secondary walls did not develop, allow an exchange of substances between adjacent living sclerenchyma cells. Mature sclerenchyma cells are unable to stretch

b-1,4 bonds. Each cellulose molecule consists of thousands of glucose subunits joined to form a flat, ribbonlike chain. From 40 to 70 of these chains lie parallel to one another and connect by hydrogen bonding to form a cellulose microfibril, a strong, tiny strand visible under the electron microscope (see Vacuole Fig. 3-10a). Nucleus Intercellular Cellulose microfibrils are cemented Cytoplasm space together by a matrix of hemicelluloses Cell wall and pectins. Hemicelluloses are a group (a) Parenchyma cells. Parenchyma cells vary in size and of polysaccharides that are more soluble Lumen structure, depending on their various functions within the than cellulose. Hemicelluloses vary in plant body. their chemical composition from one species to another. Some hemicelluThickened corner loses are composed of xyloglucan, which of cell wall consists of a backbone of b-1,4-glucose molecules to which side chains of xylose, a five-carbon sugar, are attached. DeCell wall spite their name, the chemical structure of the hemicelluloses is significantly different from that of cellulose. Pectin, another cementing polysaccharide, is less variable in its monomer composi(c) Sclerenchyma cells (fibers) in tion than are the hemicelluloses. Pectin longitudinal section (left) and cross section. Mature fibers are monomer units are a-galacturonic acid, often dead at functional maturity a six-carbon molecule that is a derivaand therefore lack nuclei and cytoplasm; the lumen is the tive of glucose. space formerly occupied by the An important component of secliving cell. ondary plant cell walls, particularly Nucleus Cytoplasm Vacuole those of wood, is lignin. Composing as (b) Collenchyma cells in longitudinal section (left) and much as 35% of the dry weight of the cross section. Note the elongated cells, evident in secondary cell wall, lignin is a strengthlongitudinal section, and the unevenly thickened cell walls, evident in cross section. ening polymer made up of monomers derived from certain amino acids. SciFIGURE 33-3 Cell types: parenchyma, collenchyma, and sclerenchyma entists have not yet determined the exact chemical structure of lignin because it is hard to remove from cellulose and or elongate. At functional maturity, when sclerenchyma tissue is other cell wall materials to which it is covalently bound. providing support for the plant body, its cells are often dead. Having discussed the four main components in plant cell walls, Sclerenchyma tissue may be located in several areas of the we can now generalize about the cell chemistry of parenchyma, plant body. Two types of sclerenchyma cells are sclereids and ficollenchyma, and sclerenchyma cells. The thin primary cell walls bers. Sclereids are cells of variable shape common in the shells of of parenchyma cells contain predominantly cellulose, although nuts and in the stones of fruits such as cherries and peaches. The they also contain hemicelluloses and pectin. Both parenchyma slightly gritty texture of pears results from the presence of clusters and collenchyma cells have primary cell walls, but their walls are of sclereids. Fibers, which are long, tapered cells that often occur chemically distinct because the thickened areas of collenchyma in patches or clumps, are particularly abundant in the wood, inner walls contain large quantities of pectin in addition to cellulose and bark, and leaf ribs (veins) of flowering plants (FIG. 33-3c). hemicelluloses. The thick secondary walls of sclerenchyma cells are chemically different because they are rich in lignin, in addition Cells of the three simple tissues vary to cellulose, hemicelluloses, and pectin. in their cell wall chemistry Parenchyma, collenchyma, and sclerenchyma cells can be distinThe vascular tissue system consists guished by the chemistry of their cell walls. Cell walls may contain of two complex tissues cellulose, hemicelluloses, pectin, and lignin. Cellulose, the most abundant polymer in the world, accounts for about 40% to 60% The vascular tissue system, which is embedded in the ground of the dry weight of plant cell walls. As discussed in Chapter 3, tissue, transports needed materials throughout the plant via two cellulose is a polysaccharide composed of glucose units joined by complex tissues: xylem and phloem (TABLE 33-3). Both xylem and

TABLE 33-3

Selected Cell Types in the Vascular Tissue System

John D. Cunningham/Visuals Unlimited

Tracheids

Pits

Fibers

James Mauseth, University of Texas

Vessel element dotted with numerous pits Perforation

Sieve plate James Mauseth, University of Texas

Vessel element Description Dead at maturity; end walls have perforations; lacks secondary wall at pits Functions Conduction of water and nutrient minerals; support Location and comments Xylem throughout plant body; vessel elements are more efficient than tracheids in conduction; shown is an LM of a longitudinal section of two vessel elements from an unidentified woody eudicot

Vessel element dotted with numerous pits

Sieve tube element

J. Robert Waaland/Biological Photo Service

Tracheid Description Dead at maturity; lacks secondary wall at pits Functions Conduction of water and nutrient minerals; support Location and comments Occurs in clumps in xylem throughout plant body; shown is an LM of a longitudinal section of tracheids from white pine (Pinus strobus) wood

Sieve tube element Description Living but lacks nucleus and other organelles at maturity; end walls are sieve plates Function Conduction of sugar in solution Location and comments Phloem throughout plant body; shown is an LM of a longitudinal section through a clump of sieve tube elements in a squash (Cucurbita) petiole (leaf stalk)

Sieve tube element

Sieve tube elements Companion cells Sieve plate 50 μm

Companion cell Description Living; has cytoplasmic connections with sieve tube element Function Assists in moving sugars into and out of sieve tube element Location and comments Phloem throughout plant body; shown is an LM of phloem from a squash (Cucurbita) petiole (leaf stalk) in cross section

phloem are continuous throughout the plant body. (Chapter 35 discusses the mechanisms of transport in xylem and phloem.) The conducting cells in xylem are tracheids and vessel elements

element into the next. A stack of vessel elements, called a vessel, resembles a miniature water pipe. Vessel elements also have pits in their side walls that permit the lateral transport (sideways movement) of water from one vessel to another.

Xylem conducts water and dissolved minerals from the roots to the stems and leaves and provides structural support. In flowering plants, xylem is a complex tissue composed of four cell types: tracheids, vessel elements, parenchyma cells, and fibers. Two of the four cell types found in xylem, the tracheids and vessel elements, conduct water and dissolved minerals. Xylem also contains fibers that provide support, and parenchyma cells, known as xylem parenchyma, that perform storage functions. Tracheids and vessel elements are highly specialized for conduction. As they develop, both types of cells undergo apoptosis, or programmed cell death. As a result, mature tracheids and vessel elements are dead and therefore hollow; only their cell walls remain. Tracheids, the chief water-conducting cells in gymnosperms (such as pine) and seedless vascular plants (such as ferns), are long, tapering cells located in patches or clumps (FIG. 33-4a). Water is conducted upward, from roots to shoots, passing from one tracheid into another through wide pits. In addition to tracheids, flowering plants possess extremely efficient vessel elements (FIG. 33-4b). The cell diameters of vessel elements are usually greater than those of tracheids. Vessel elements are hollow, but unlike tracheids, the end walls have holes, known as perforations, or are absent. Vessel elements are stacked one on top of the other, and water is conducted readily from one vessel

Sieve tube elements are the conducting cells of phloem Phloem conducts food materials, that is, carbohydrates formed in photosynthesis, throughout the plant and provides structural support. In flowering plants, phloem is a complex tissue composed of four cell types: sieve tube elements, companion cells, fibers, and phloem parenchyma cells (FIGS. 33-4c and 33-4d). Fibers, which are usually extensive in the phloem of herbaceous plants, provide additional structural support for the plant body. Food materials are conducted in solution—that is, dissolved in water—through the sieve tube elements, which are among the most specialized plant cells. Sieve tube elements are joined end to end to form long sieve tubes. The cells’ end walls, called sieve plates, have a series of holes through which cytoplasm extends from one sieve tube element into the next. Sieve tube elements are living at maturity, but many of their organelles, including the nucleus, vacuole, mitochondria, and ribosomes, disintegrate or shrink as they mature. Sieve tube elements are among the few eukaryotic cells that can function without nuclei. These cells typically live for less than a year. There are, however, notable exceptions. Certain palms have sieve tube elements that remain alive for approximately 100 years. Adjacent to each sieve tube element is a companion cell that assists in the functioning of the sieve tube element. The compan-

Xylem

Phloem

End wall with perforations

Sieve plate with pores Sieve tube element Phloem parenchyma cells

Pits Cell wall Lumen

Lateral sieve area Plasmodesma Companion cell

(a) Tracheid (xylem cell). The tracheid is opened to reveal the cell’s appearance in cross section. At maturity, tracheids are usually dead.

(b) Vessel element (xylem cell). The end walls of vessel elements are perforated. Vessel elements are joined end to end, from roots to leaves and other shoot parts.

(c) Sieve tube element, showing the sieve plate. Sieve tube elements are joined end to end to form sieve tubes.

(d) Longitudinal section of phloem tissue. Sieve tube elements, companion cells, and phloem parenchyma cells. Fiber cells are not shown.

ion cell is a living cell, complete with a nucleus. This nucleus is thought to direct the activities of both the companion cell and sieve tube element. Numerous plasmodesmata—cytoplasmic connections through which cytoplasm extends from one cell to another (see Chapter 5)—link a companion cell with its adjoining sieve tube element. The companion cells play an essential role in moving sugar into the sieve tube elements for transport to other parts of the plant. TABLE 33-4

Epidermal cell Description Relatively unspecialized cell with thin primary wall; outer wall often thicker and covered by a noncellular waxy layer (cuticle) Functions Protective covering over surface of plant body; helps reduce water loss Location and comments Epidermis is usually one cell thick; shown is an LM of a cross section through epidermis of an ivy (Hedera helix) stem

James Mauseth, University of Texas

Epidermis

Collenchyma

Guard cell Description Chloroplast-containing cell that occurs in pairs; pair changes shape to open and close stomatal pore Function Opens and closes stomatal pore Location and comments Epidermis of stems and leaves; shown is an LM of the epidermis of a spiderwort (Tradescantia virginiana) leaf

Guard cell Stomatal pore Epidermal cell Dwight R. Kuhn

The dermal tissue system—the epidermis and periderm—provides a protective covering over plant parts (TABLE 33-4). In herbaceous plants, the dermal tissue system is a single layer of cells called the epidermis. Woody plants initially produce an epidermis, but it splits apart as a result of the production of additional woody

Selected Cell Types in the Dermal Tissue System Cuticle

Guard cell

Trichome Description Hair or other epidermal outgrowth; may be unicellular or multicellular; occurs in variety of sizes and shapes Functions Varied; absorption; secretion; excretion; protection; reduction of water loss Location and comments Epidermis; shown is an SEM of a nettle (Solanum carolinense) leaf, which has trichomes that break off inside the skin of animals and inject irritating substances that cause a stinging sensation

Trichomes

Biophoto Associates

Epidermis

Cork cells Cork cambium Cork parenchyma

Periderm

Remnants of epidermis

Cengage

The dermal tissue system consists of two complex tissues

Cork cell Description Dead at maturity; cell walls impregnated with waterproof material (suberin) Functions Reduces water loss and prevents disease-causing organisms from penetrating Location and comments Produced in large numbers; cork often forms just under the epidermis; replaces epidermis in older stems and roots; shown is an LM of a cross section through periderm of a geranium (Pelargonium) stem

tissues inside the epidermis as the plant increases in girth. Periderm, which replaces the epidermis in the stems and roots of older woody plants, composes the outer bark. Epidermis is the outermost layer of a herbaceous plant

The epidermis is a complex tissue composed primarily of relatively unspecialized living cells. Dispersed among these cells are more specialized guard cells and outgrowths called trichomes (discussed later). In most plants, the epidermis consists of a single layer of flattened cells (FIG. 33-5). Epidermal cells generally contain no chloroplasts and are therefore transparent, so light can penetrate the interior tissues of stems and leaves. In both stems and leaves, photosynthetic tissues lie beneath the epidermis. An important requirement of the aerial parts (stems, leaves, flowers, and fruits) of a plant is the ability to control water loss. Epidermal cells of aerial parts secrete a waxy cuticle over the surface of their exterior walls; this waxy layer greatly restricts water loss from plant surfaces. Although the cuticle is extremely efficient at preventing most water loss through epidermal cells, it also slows the diffusion of CO2 , needed for photosynthesis, from the atmosphere into the leaf or stem. The diffusion of CO2 is facilitated by stomata (sing.,

Cell wall Cytoplasm Vacuole Nucleus

stoma). Stomata are minute pores in the epidermis that are surrounded by two cells called guard cells (see Fig. 34-5). Many gases, including CO2, oxygen, and water vapor, pass through the stomata by diffusion. Stomata are generally open during the day when photosynthesis is occurring, and the water loss that also takes place when stomata are open provides some evaporative cooling. At night, stomata usually close. In drought conditions, the need to conserve water overrides the need to cool the leaves and exchange gases. Thus, during a drought, the stomata close in the daytime. Stomata are discussed in greater detail in Chapter 34. The epidermis also contains special outgrowths, or hairs, called trichomes, which occur in many sizes and shapes and have a variety of functions. Plants that tolerate salty environments such as the seashore often have specialized trichomes on their leaves to remove excess salt that accumulates in the plant. The presence of trichomes on the aerial parts of desert plants may increase the reflection of light off the plants, thereby cooling the internal tissues and decreasing water loss. Other trichomes have a protective function. For example, the trichomes on stinging nettle leaves and stems contain irritating substances that may discourage herbivorous animals from eating the plant. Root hairs are simple, unbranched trichomes that increase the surface area of the root epidermis (which comes into contact with the soil) for more effective water and mineral absorption. Periderm replaces epidermis in woody plants Periderm, a tissue that can be anywhere from several to many cell layers thick, forms under the epidermis to provide a new protective covering as the epidermis is destroyed. As a woody plant continues to increase in girth, its epidermis is sloughed off, exposing the periderm. Periderm forms the outer bark of older stems and roots. It is a complex tissue composed mainly of cork cells and cork parenchyma cells. Cork cells are dead at maturity, and their walls are heavily coated with a waterproof substance called suberin, which helps reduce water loss. Cork parenchyma cells function primarily in storage. Periderm is discussed further in Chapters 35 and 36.

Review ■ ■

■

■

■

What are some of the functions of roots? of shoots? What are the three tissue systems in plants? Describe the functions of each. How do parenchyma, collenchyma, and sclerenchyma tissues differ in cell structure and function? What are the functions of xylem and phloem? Describe the conducting cells that occur in each of these complex tissues. How are epidermis and periderm alike? How are they different?

33.2 PLANT MERISTEMS ■ ■ LEARNING OBJECTIVES FIGURE 33-5 Cell types: epidermal cells Epidermal cells from three different plants. Note that regardless of the cell shape, epidermal cells fit tightly together.

5 State how growth in plants differs from growth in animals. 6 Distinguish between primary and secondary growth. 7 Distinguish between apical meristems and lateral meristems.

Primary growth takes place at apical meristems Primary growth occurs as a result of the activity of apical meristems, areas located at the tips of roots and shoots, including within the buds of stems. Buds are dormant embryonic shoots that eventually develop into branches. Primary growth is evident when a root tip is examined (FIG. 33-6). A protective layer of cells called a root cap covers the root tip. Directly behind the root cap is the root apical meristem, which consists of meristematic cells. Meristematic cells, which are cube shaped, remain small because they are continually dividing. (In meristems, as daughter cells begin to enlarge, one or both will divide again.) Farther from the root tip, just behind the area of cell division, is an area of cell elongation where the cells have been displaced from the meristem. Here the cells are no longer dividing but instead are growing longer, pushing the root tip ahead of them, deeper into the soil. Some differentiation also begins to occur in the area of cell elongation, and immature tissues such as differentiating xylem and phloem become evident. The tissue systems continue to develop and differentiate into primary tissues (epidermis, xylem, phloem, and ground tissues) of the adult plant. Farther from the tip, behind the area of cell elongation, most cells have completely differentiated and are fully mature. Root hairs are evident in this area.

Branch root

Root hairs

Area of cell maturation

Area of cell elongation Root cap

Apical meristem (area of cell division)

Cengage

One difference between plants and animals is the location of growth. When plants grow, their cells divide only in specific areas, called meristems, which are composed of cells whose primary function is to form new cells by mitotic division. (In contrast, when a young animal is growing, all parts of its body grow, although not necessarily at the same rate.) Meristematic cells, also known as stem cells, do not differentiate. Instead, they retain the ability to divide by mitosis, a trait that many differentiated cells lose. The persistence of mitotically active meristems means that plants, unlike most animals, can grow throughout their entire lifespan. This ability of roots and stems to grow throughout a plant’s life is known as indeterminate growth. In contrast, many leaves and flowers have determinate growth; that is, they stop growing after reaching a certain size. The size of leaves and flowers with determinate growth varies from species to species and from individual to individual, depending on the plant’s genetic programming and on environmental conditions such as availability of sunlight, water, and essential minerals. Two kinds of meristematic growth may occur in plants. Primary growth is an increase in stem and root length. All plants have primary growth, which produces the entire plant body in herbaceous plants and the young, soft shoots and roots in woody trees and shrubs. Secondary growth is an increase in the girth of a plant. For the most part, only gymnosperms and woody eudicots have secondary growth. (Woody eudicots include trees and shrubs such as oak, sycamore, ash, cherry, apple, beech, and maple.) Tissues produced by secondary growth comprise the wood and bark, which make up most of the bulk of trees and shrubs. A few annuals—geranium and sunflower, for example—undergo limited secondary growth despite the fact that they lack obvious wood and bark tissues.

FIGURE 33-6 Animated A root tip The root apical meristem (where cells divide and thus increase in number) is protected by a root cap. Farther from the tip is an area of cell elongation where cells enlarge and begin to differentiate. The area of cell maturation has fully mature, differentiated cells. Note the young root hairs in this area and on the root of a young radish (Raphanus sativus) seedling, which is approximately 5 cm (2 in) long (left).

A shoot apex (such as the terminal bud) is quite different in appearance from a root tip (FIG. 33-7). A dome of minute, regularly arranged meristematic cells—the shoot apical meristem—is located in each shoot apex. Leaf primordia (developing leaves) and bud primordia (developing buds) arise from the shoot apical meristem. The tiny leaf primordia cover and protect the shoot apical meristem. As the cells formed by the shoot apical meristem elongate, the shoot apical meristem is pushed upward. Subsequent cell divisions produce additional stem tissue and cause new leaf and bud primordia to appear. Farther from the stem tip, the immature cells differentiate into the three tissue systems of the mature plant body.

Secondary growth takes place at lateral meristems Trees and shrubs undergo both primary and secondary growth. These plants increase in length by primary growth and increase in girth by secondary growth. The increase in girth, which occurs in areas that are no longer elongating, is due to cell divisions that take place in lateral meristems, areas extending along the entire length of the stems and roots except at the tips. Two lateral meristems, the vascular cambium and the cork cambium, are responsible for secondary growth, which is the formation of secondary tissues: secondary xylem, secondary phloem, and periderm (FIG. 33-8). The vascular cambium is a layer of meristematic cells that forms a long, thin, continuous cylinder within the stem and root.

Older leaf Leaf primordia Apical meristem James Mauseth, University of Texas

Coleus

Older leaf Trichome Bud primordium 100 μm

FIGURE 33-7 A shoot apex LM of a longitudinal section through a shoot apex of Coleus, showing the shoot apical meristem, leaf primordia, and bud primordia.

It is located between the wood and bark of a woody plant. Division of cells of the vascular cambium adds more cells to the wood (secondary xylem) and inner bark (secondary phloem). The cork cambium, located in the outer bark, is composed of a thin cylinder or irregular arrangement of meristematic cells. Cells of the cork cambium divide and form cork cells toward the outside and one or more underlying layers of cork parenchyma cells that function in storage. Collectively, cork cells, cork cambium, and cork parenchyma make up the periderm. We are now ready to give a more precise definition of bark. Bark, the outermost covering over woody stems and roots, consists of all plant tissues located outside the vascular cambium.

Bark has two regions, a living inner bark composed of secondary phloem and a mostly dead outer bark composed of periderm. Chapters 35 and 36 present a more comprehensive discussion of secondary growth.

Review ■

■ ■

What is the role of plant meristems? Does animal growth involve meristems? What is primary growth? secondary growth? What are apical meristems? lateral meristems?

33.3 DEVELOPMENT OF FORM Outer bark (periderm) Inner bark (secondary phloem)

■ ■ LEARNING OBJECTIVES 8 Distinguish between cell division and cell expansion, and describe the roles of the preprophase band and cellulose microfibrils.

9 Describe the relationship between cell determination and cell differentiation, and between pattern formation and morphogenesis.

10 Explain why the model organism Arabidopsis is so useful in the study of plant development.

Bark

Wood (secondary xylem) Vascular cambium

FIGURE 33-8 Secondary growth The vascular cambium, a thin layer of cells sandwiched between the wood and bark, produces the secondary vascular tissues: the wood, which is secondary xylem, and the inner bark, which is secondary phloem. The cork cambium produces the periderm, the outer bark tissue that replaces the epidermis in a plant with secondary growth.

Plants are growing, dynamic organisms. Development in plants encompasses all the changes that take place during the entire life of an individual. Of particular interest in developmental biology is the process by which cells specialize and organize into a complex organism. For example, how does a microscopic, unicellular zygote give rise to a multicellular embryo in a seed? (See the stages of embryonic development in a typical eudicot in Figure 37-8.) How does the embryo in the seed give rise to the leaves, stems, and roots of a juvenile plant, and what mechanisms underlie the transition from a nonreproductive juvenile plant to an adult plant capable of flowering (FIG. 33-9)? We now examine several aspects of plant development that affect form in plants. Flowering is discussed elsewhere in this book (see the ABC model in Chapter 17).

250 mm

Leafy shoot derives from shoot apical meristem

Shoot apical meristem

5 mm Root system derives from root apical meristem

Root apical meristem

(a) Seedling

(b) Juvenile plant (nonreproductive)

(c) Adult plant (reproductive)

FIGURE 33-9 Growth of an Arabidopsis seedling into an adult plant Almost all of the differentiated tissues formed after a seed is germinated (often referred to as postembryonic growth) come from a small number of stem cells in the root and shoot apical meristems.

The plane and symmetry of cell division affect plant form

determines the plane in which cells divide, and there is no preprophase band.

You have seen that plant growth is localized into meristems for cell division. At these sites, the cells undergo cell division in an orderly manner. In the onion root apical meristem, for example, files of cells can be traced back to the meristem region (FIG. 33-10). These cells divided in the same plane to produce each file of cells. Cell divisions in one plane, two planes, or more help to establish the tissue patterns that will be produced during development. How is the plane of cell division for a particular meristematic cell determined? In plant cells, a dense array of microtubules forms a preprophase band just inside of the plasma membrane, encircling the nucleus much like a belt. (Chapter 4 contains a discussion of the dynamic nature of microtubules.) The preprophase band appears just prior to mitosis and determines the plane in which the cell will divide (FIG. 33-11). The preprophase band, which is located where the cell plate will form following mitosis and cytokinesis, also helps to orient the mitotic spindle. The preprophase band is a plant characteristic; in animal cells, the orientation of the spindle

The orientation of cellulose microfibrils affects the direction of cell expansion In plants, growth occurs by both an increase in the number of cells and an increase in cell size. Cell division and cell expansion (also called elongation) are governed by a genetic program that interacts with environmental signals from both the external and internal environments. In growth, a newly formed plant cell absorbs water into its vacuole by osmosis. The water increases the cell’s turgor pressure, hydrostatic pressure that develops within a walled cell and presses outward against the plasma membrane. Turgor pressure provides a force that exerts pressure on the cell wall and causes it to expand. In an onion root cell, the vacuole increases in size by some 30 to 150 times during elongation. In growing tissue, the arrangement of cellulose microfibrils in the wall prevents the cell from expanding equally in all directions

3

Cells form a single file

2

Two cells become three

Maximum rate of cell division Meristematic zone

Cell division in the same plane as before

One cell becomes two

Plane of cell division

© Visuals Unlimited / Corbis

1

Root cap

200 μm

(a) Cells that divide in the same plane form a row, or file, of cells.

(b) A single file of cells in the onion root tip is labeled. If you look closely, you can see many other files.

FIGURE 33-10 The plane of cell division and the onion root tip The root apical meristem maintains postembryonic root growth.

due to turgor pressure. When the microfibrils in the cell wall are oriented in the same direction, the wall expands perpendicularly to them (FIG. 33-12).

Cell differentiation depends in part on a cell’s location By themselves cell division and cell expansion would produce only a formless heap of similar cells. The development of plant form is the result of a balanced combination of several fundamental processes that go beyond cell division and cell expansion: cell differentiation and cell determination, and pattern formation and morphogenesis.

As growth proceeds, cell division gives rise to increasing numbers of cells, which serve as the building blocks of development. At various times certain cells become biochemically and structurally specialized to carry out specific functions through a process known as cell differentiation. Generally, a differentiated animal cell is irreversibly committed to its fate, but that is not the case with plants. As discussed earlier in the chapter, parenchyma cells can differentiate into other kinds of cells, for example, when the plant body is injured. Cell differentiation occurs through cell determination, a series of molecular events in which the activities of certain genes are altered in ways that cause a cell to progressively commit to a particular differentiation pathway. As cell determination proceeds, a

Cellulose microfibrils

Nucleus

Coalescing vacuoles

Elongated, enlarged vacuole

FIGURE 33-12 Animated Direction of cell expansion The preprophase band encircles the nucleus and indicates the plane of an upcoming cell division.

FIGURE 33-11 Preprophase band

young cell’s fate becomes more and more limited, restricting its development to a limited set of tissue types. During cell differentiation, a cell’s physical appearance may or may not change significantly. Nevertheless, when cell determination is complete, a cell becomes structurally and functionally recognizable—as an epidermal cell, for example—and its pattern of gene activity differs from that of other cell types. Thus, differentiated cells of the plant body have differential gene expression— that is, different genes are active in different cells at the same time. Differential gene expression is responsible for variations in chemistry, behavior, and structure among cells. Tissue cultures involve differential gene expression Cells can be isolated from certain plants and grown in a chemically defined, sterile nutrient medium. In initial experiments with such cultures, plant cells could be kept alive, but they did not divide. Researchers later discovered that adding certain natural materials, such as the liquid endosperm of coconut, also known as coconut milk, induced cells to divide in culture. By the late 1950s, plant cells from a variety of sources could be cultured successfully, dividing to produce a mass of disorganized, relatively undifferentiated cells, or callus. In 1958, F. C. Steward, a plant physiologist at Cornell University, succeeded in generating an entire carrot plant from a single callus cell derived from a carrot root (see Fig. 17-2). This demonstrated that each plant cell contains a genetic blueprint for all features of an entire organism. His work also showed that an entire plant can be grown from a single cell, provided differential gene expression occurs so that the proper genes are expressed at the appropriate times. Since Steward’s pioneering work, biologists have successfully cultured many plants by using a variety of cell sources. Plants have been regenerated from different tissues, organ explants (excised organs or parts such as root apical meristems), and single cells.

Following cell division, cellulose microtubules are deposited into new cell walls in an orderly fashion that permits cell expansion. The orientation of the cellulose microfibrils determines the direction of cell expansion. Note that most of the cellulose microfilbrils are oriented transversely in this figure, which results in longitudinal cell expansion.

Cell and tissue culture techniques help answer many fundamental questions involving growth and development in plants. These techniques also have great practical potential. Using tissue culture, researchers can regenerate many genetically identical plants from the cells of a single, genetically superior plant. Many kinds of plants—from orchids and African violets to coastal redwoods— have been cultured in this way.

Morphogenesis occurs through pattern formation Cell differentiation by itself does not explain overall plant form. The differentiated cells must become progressively organized, shaping the intricate pattern of tissues and organs that characterizes a multicellular plant. This development of form, in which differentiated cells in specific locations become spatially organized into recognizable structures, is known as morphogenesis. Morphogenesis proceeds through the process of pattern formation, a series of steps requiring signaling between cells, changes in the shapes and metabolism of certain cells, and precise cell interactions. Pattern formation organizes cells into threedimensional structures (FIG. 33-13). Depending on their location, cells are exposed to different concentrations of signaling molecules that specify positional information—that is, where the cell is located relative to the body’s axes. This information affects cell differentiation and tissue formation. Thus, where a given cell is located often determines what it will become when it matures. Arabidopsis mutants provide crucial insights into plant development Until the late 1970s, biologists knew little about how genes interact to control development. However, rapid progress in molecular biology led scientists to search for developmental mutants and to

(a) Examples of leaf shapes.

(b) Examples of leaf margins.

FIGURE 33-13 Leaf variation and pattern formation

apply sophisticated techniques to study them. (A mutant is an individual with an abnormal phenotype caused by a gene mutation.) Biologists are now studying how genes are activated, inactivated, and modified to control development. Eventually, scientists expect to reach a major goal of systems biology—understanding how a single cell (a fertilized egg) develops into a multicellular plant. You may recall from Chapter 17 that mouse-ear cress (Arabidopsis thaliana) is the organism most widely used to study the genetic control of development in plants. When biologists insert cloned foreign genes into Arabidopsis cells, the genes integrate into the chromosomes and are expressed. The researchers then induce these transformed cells to differentiate into transgenic plants. Botanists use chemical mutagens to produce mutant strains and have isolated many developmental mutants of Arabidopsis. Consider the monopteros (mp) gene, which codes for a transcription factor affecting apical-basal pattern formation and the development of vascular tissues. Mutant mp seedlings lack basal structures, including a primary root (FIG. 33-14). Such mutant phenotypes have helped scientists identify the genes essential for normal development. Arabidopsis has also had its relatively small genome—about 26,700 genes—sequenced. The success of the Arabidopsis sequencing project led to an international initiative by plant biologists to understand the functions of all the genes in the Arabidopsis genome (TABLE 33-5). This functional genomic information will lead to a far deeper understanding of plant development and evolutionary history.

T. Berleth/ G. Jurgens

Leaves are quite variable in form—there are dozens of different leaf shapes and margins. Leaves on juvenile plants are often shaped differently than those on adult plants, and some adult plants have several leaf shapes on the same individual. Pattern formation, which is under genetic control, is largely responsible for leaf variation. (Also see Figure 34-2 for more examples of variation in leaves.)

FIGURE 33-14 Monopteros mutation in Arabidopsis seedling The monopteros gene affects development of the root system. The wild type seedling (left) has a normal monopteros gene, whereas the mutant (right) lacks basal structures.

Review ■

■ ■ ■

Which process typically occurs first, cell differentiation or cell expansion? What are some kinds of events that contribute to pattern formation? How does pattern formation lead to morphogenesis? Why is Arabidopsis such a useful model organism for plant development?

Known Gene Functions in the Arabidopsis Genome

Function

Percent of Genome

DNA/RNA metabolism Signal transduction Development Energy metabolism Cell division Transport Transcription Response to environment Protein metabolism Other biological processes Other cellular processes Other metabolic functions Unknown functions

1 2 2 3 3 4 4 4 7 11 17 18 24

© Nigel Cattlin/Alamy

TABLE 33-5

Source: The Arabidopsis Information Resource, 2007.

■ ■

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

33.1 (page 709) 1 Discuss the functions of various parts of the vascular plant body, including the nutrient- and water-absorbing root system and the photosynthesizing shoot system. ■ The vascular plant body typically consists of a root system and a shoot system. The root system is generally underground and obtains water and dissolved minerals for the plant. Roots also anchor the plant firmly in place. ■ The shoot system is generally aerial and obtains sunlight and exchanges gases such as CO2, oxygen, and water vapor. The shoot system consists of a vertical stem that bears leaves and reproductive structures. Buds, undeveloped embryonic shoots, develop on stems.

2 Describe the structure and functions of the ground tissue system (parenchyma tissue, collenchyma tissue, and sclerenchyma tissue). ■ The ground tissue system consists of three tissues with a variety of functions. Parenchyma tissue is composed of living parenchyma cells that have thin primary cell walls. Functions of parenchyma tissue include photosynthesis, storage, and secretion. ■ Collenchyma tissue consists of collenchyma cells with unevenly thickened primary cell walls. This tissue provides flexible structural support. ■ Sclerenchyma tissue is composed of sclerenchyma cells—sclereids or fibers—that have both primary cell walls and secondary cell walls. Sclerenchyma cells are often dead at maturity but provide structural support. Learn more about parenchyma, collenchyma, and sclerenchyma cells by clicking on the figure in CengageNOW.

3 Describe the structure and functions of the vascular tissue system (xylem and phloem). ■ The vascular tissue system conducts materials throughout the plant body and provides strength and support. ■ Xylem is a complex tissue that conducts water and dissolved minerals. The actual conducting cells of xylem are tracheids and vessel elements. ■ Phloem is a complex tissue that conducts sugar in solution. Sieve tube elements are the conducting cells of phloem; they are assisted by companion cells. Learn more about xylem and phloem by clicking on the figure in CengageNOW.

4 Describe the structure and functions of the dermal tissue system (epi-

Learn more about the plant body by clicking on the figure in CengageNOW.

dermis and periderm). The dermal tissue system is the outer protective covering of the ■ plant body.

■

■

The epidermis is a complex tissue that covers the herbaceous plant body. The epidermis that covers aerial parts secretes a waxy cuticle that reduces water loss. Stomata permit gas exchange between the interior of the shoot system and the surrounding atmosphere. Trichomes are outgrowths, or hairs, that occur in many sizes and shapes and have a variety of functions. The periderm is a complex tissue that covers the plant body in woody plants.

33.2 (page 718) 5 State how growth in plants differs from growth in animals. Growth in plants, unlike that in animals, is localized in specific regions called meristems. Meristems are composed of cells whose primary function is to form new cells by mitotic division. 6 Distinguish between primary and secondary growth. ■ Primary growth is an increase in stem or root length. Primary growth occurs in all plants. ■

See root and shoot development by clicking on the figures in CengageNOW. ■

Secondary growth is an increase in stem or root girth (thickness). Secondary growth typically occurs in long cylinders of meristematic cells throughout the length of older stems and roots. Learn more about secondary growth by clicking on the figures in CengageNOW.

7 Distinguish between apical meristems and lateral meristems. ■

■

Primary growth results from the activity of apical meristems that are localized at the tips of roots and shoots and within the buds of stems. The two lateral meristems responsible for secondary growth are the vascular cambium and the cork cambium.

Growth occurs by both cell division (an increase in the number of cells) and cell expansion (an increase in cell size). ■ In plant cells, a dense array of microtubules forms a preprophase band just inside of the plasma membrane. The preprophase band appears just prior to mitosis and determines the plane in which the cell will divide. Absorption of water into a cell’s vacuole increases the turgor ■ pressure, which exerts pressure on the cell wall. The arrangement of cellulose microfibrils in the wall prevents the wall from expanding equally in all directions. When microfibrils are oriented the same way, the wall expands perpendicular to the microfibrillar orientation. 9 Describe the relationship between cell determination and cell differentiation, and between pattern formation and morphogenesis. ■ In cell differentiation, cells become biochemically and structurally specialized to carry out specific functions. Cell differentiation occurs through cell determination, a series of molecular events in which the activities of certain genes are altered in ways that cause a cell to progressively commit to a particular differentiation pathway. ■ The development of form, in which differentiated cells in specific locations become spatially organized into recognizable structures, is known as morphogenesis. Morphogenesis proceeds through the process of pattern formation, a series of steps requiring signaling between cells, changes in the shapes and metabolism of certain cells, and precise cell interactions. 10 Explain why the model organism Arabidopsis is so useful in the study of plant development. ■ Botanists have isolated many developmental mutants of Arabidopsis. Such mutant phenotypes have helped scientists identify genes essential for normal development. The genome of Arabidopsis has been sequenced, and plant biologists are currently determining the functions of all of this plant’s genes. ■

33.3 (page 720) 8 Distinguish between cell division and cell expansion, and describe the roles of the preprophase band and cellulose microfibrils.

T E S T YO U R U N D E R S TA N D I N G 1. Most of the plant body consists of the _____ tissue system. (a) ground (b) vascular (c) periderm (d) dermal (e) cortex

5. Sclereids and fibers are examples of which plant tissue? (a) parenchyma (b) collenchyma (c) sclerenchyma (d) xylem (e) epidermis

2. The cell walls of parenchyma cells (a) contain large quantities of pectin in the thickened corners (b) are rich in lignin but do not contain hemicelluloses and pectin (c) are predominantly cellulose, although they also contain hemicelluloses and pectin (d) contain cellulose, hemicelluloses, and lignin in approximately equal amounts (e) contain hemicelluloses, pectin, and lignin but no cellulose

6. Which of the following statements about the vascular tissue system is not true? (a) xylem and phloem are continuous throughout the plant body (b) xylem not only conducts water and dissolved minerals but also provides support (c) four cell types occur in phloem: sieve tube elements, companion cells, tracheids, and vessel elements (d) sieve tube elements lack nuclei (e) vessel elements are hollow, and their end walls have perforations or are entirely dissolved away

3. Which tissue system provides a covering for the plant body? (a) ground (b) vascular (c) periderm (d) dermal (e) cortex 4. The two simple tissues that are specialized for support are (a) parenchyma and collenchyma (b) collenchyma and sclerenchyma (c) sclerenchyma and parenchyma (d) parenchyma and xylem (e) xylem and phloem

7. Minute pores known as _____ dot the surface of the epidermis of leaves and stems; each pore is bordered by two _____. (a) stomata; guard cells (b) stomata; fibers (c) sieve tube elements; companion cells (d) sclereids; guard cells (e) cuticle; guard cells

8. Primary growth, an increase in the length of a plant, occurs at the (a) cork cambium (b) apical meristem (c) vascular cambium (d) lateral meristem (e) periderm

12. Label the three tissue systems in the plant body.

9. The preprophase band (a) appears just prior to mitosis in both plant and animal cells (b) determines the plane in which the cell will divide (c) attaches to cellulose microfibrils in the cell wall (d) is an essential part of pattern formation in plants (e) increases a cell’s turgor pressure 10. Cell differentiation occurs through _____, whereas morphogenesis involves _____. (a) pattern formation; cell determination (b) cell division; cell expansion (c) cell expansion; pattern formation (d) cell determination; cell division (e) cell determination; pattern formation 11. The monopteros mutant (a) lacks a primary root (b) develops normally until the plant becomes reproductive (c) has abnormal leaf morphogenesis (d) was discovered in rice after its genome was sequenced (e) is one of only two developmental mutants known in Arabidopsis.

(a) Leaf

(b) Stem

(c) Root

CRITICAL THINKING 1. Grasses have a special meristem situated at the base of the leaves. Relate this information to what you know about the growth of grass after you mow the lawn.

than flowering plants, have tracheids only. Do these differences in xylem structure help explain the difference in the success of these two plant groups? Explain your answer.

2. A couple carved a heart with their initials into a tree trunk 4 ft above ground level; the tree was 25 ft tall at the time. Twenty years later the tree was 50 ft tall. How far above the ground were the initials? Explain your answer.

5. Plant biologists are currently working to identify those Arabidopsis genes with unknown functions (see Table 33-5). If you were assigned to determine the function of a particular gene, how might you approach this task?

3. Sclerenchyma in plants is the functional equivalent of bone in humans (both sclerenchyma and bone provide support). However, sclerenchyma is dead, whereas bone is living tissue. What are some of the advantages of a plant having dead support cells? Can you think of any disadvantages? Explain your answer.

6. SCIENCE, TECHNOLOGY, AND SOCIETY. Why is knowledge gained from sequencing the Arabidopsis genome important even though Arabidopsis has no commercial value?

4. EVOLUTION LINK. Flowering plants have both tracheids and vessel elements in their xylem. Conifers, which evolved earlier

Additional questions are available in CengageNOW at www.cengage.com/ login.

34

Leaf Structure and Function

Andrew Syred/Science Photo Library

Colorized SEM of the interior of a spinach leaf. The loose arrangement of internal cells in spinach (Spinacia oleracea) optimizes the diffusion of necessary raw materials such as carbon dioxide and water into photosynthetic cells.

KEY CONCEPTS

P

lants allocate many resources into the production of leaves that are often short-lived. Each year, a large maple tree, for example, may

34.1 Leaf structure reflects its primary function of

produce 47 m2 (500 ft2) of leaves, which may weigh more than 113 kg

photosynthesis.

(250 lb). The metabolic cost of producing so many leaves is high, but

34.2 Opening and closing of stomata affect carbon dioxide availability in the daily cycle of photosynthesis.

necessary. Simply put, leaves are essential to the tree’s survival, so plants

34.3 Transpiration, the loss of water vapor from leaf surfaces, keeps leaves from getting overheated and promotes water transport in the plant. Guttation is the exudation of liquid water from leaf margins.

must invest in their production. In most vascular plants, leaves are the primary organs to effectively collect solar energy and convert it into chemical energy. That is, leaves gather the sunlight necessary for photosynthesis, the biological process

34.4 Leaf abscission is the seasonal removal of deciduous

that converts radiant energy into the chemical energy of carbohydrate

leaves.

molecules. Plants use these molecules as starting materials to synthesize

34.5 Some leaves are modified for special functions in addition to photosynthesis and transpiration.

all other organic compounds and as fuel to provide energy for metabolism. During a single summer, the leaves of a maple tree will fix about 454 kg (1000 lb) of carbon dioxide (CO2) into organic compounds. The structure of a leaf is superbly adapted for its primary function of photosynthesis. Most leaves are thin and flat, a shape that allows optimal absorption of light energy and the efficient internal diffusion of gases such as CO2 and O2. As a result of their ordered arrangement on the stem,

leaves efficiently catch the sun’s rays with a minimum of “interference” from neighboring leaves. The leaves form an intricate green mosaic, bathed in sunlight and atmospheric gases. To control water loss, a thin, transparent layer of wax covers

Blade

the leaf surface. Such structural adaptations are compromises between competing needs, and some features that optimize photo-

Veins

synthesis promote water loss. For example, plants have minute pores that allow gas exchange for photosynthesis, but these openings also let water vapor escape into the atmosphere. Thus, leaf structure represents a trade-off between photosynthesis and water conservation.

Petiole

In this chapter we examine leaf form and function. Leaf form is highly variable, and thousands of different leaf shapes have evolved over millions of years. Despite this diversity, most leaves “work” in Axillary bud

the same way, and their internal structures (see photograph) support this similarity in function. Stipules

Stem

34.1 LEAF FORM AND STRUCTURE ■ ■ LEARNING OBJECTIVES 1 2

3 4

Discuss variation in leaf form, including simple versus compound leaves, leaf arrangement on the stem, and venation patterns. Describe the major tissues of the leaf (epidermis, photosynthetic ground tissue, xylem, and phloem) and label them on a diagram of a leaf cross section. Compare leaf anatomy in eudicots and monocots. Relate leaf structure to its function of photosynthesis.

Foliage leaves are the most variable of plant organs, so much so that plant biologists have developed specific terminology to describe their shapes, margins (edges), vein patterns, and the way they attach to stems. Because each leaf is characteristic of the species on which it grows, many plants can be identified by their leaves alone. Leaves may be round, needlelike, scalelike, cylindrical, heart-shaped, fanshaped, or thin and narrow. They vary in size from those of the raffia palm (Raphia ruffia), whose leaves often grow more than 20 m (65 ft) long, to those of water-meal (Wolffia), whose leaves are so small that 16 of them laid end to end measure only 2.5 cm (1 in). The broad, flat portion of a leaf is the blade; the stalk that attaches the blade to the stem is the petiole. Some leaves also have stipules, which are leaflike outgrowths usually present in pairs at the base of the petiole (FIG. 34-1). Some leaves do not have petioles (that is, the blades are attached directly to the stem) or stipules; leaves without a petiole are said to be sessile. Leaves may be simple (having a single blade) or compound (having a blade divided into two or more leaflets) (FIG. 34-2a). Sometimes it is difficult to tell whether a plant has formed one compound leaf or a small stem bearing several simple leaves. One easy way to determine if a plant has simple or compound leaves is to look for axillary buds, so-called because each develops in a leaf axil (the angle between the stem and petiole). Axillary buds form at the base of a leaf, whether it is simple or compound. However, axillary buds never develop at the base of leaflets. Also, the leaflets

FIGURE 34-1 Parts of a leaf A geranium leaf consists of a blade, a petiole, and two stipules at the base of the leaf. Note the axillary bud in the leaf axil.

of a compound leaf lie in a single plane (you can lay a compound leaf flat on a table), whereas simple leaves usually are not arranged in one plane on a stem. Leaves are arranged on a stem in one of three possible ways (FIG. 34-2b). Plants such as beeches and walnuts have an alternate leaf arrangement, with one leaf at each node, the area of the stem where one or more leaves are attached. In an opposite leaf arrangement, as occurs in maples and ashes, two leaves grow at each node. In a whorled leaf arrangement, as in catalpa trees, three or more leaves grow at each node. Leaf blades may possess parallel venation, in which the primary veins—strands of vascular tissue—run approximately parallel to one another (generally characteristic of monocots), or netted venation, in which veins are branched in such a way that they resemble a net (generally characteristic of eudicots; FIG. 34-2c).1 Netted veins can be pinnately netted, with major veins branching off in succession along the entire length of the midvein (main or central vein of a leaf), or palmately netted, with several major veins radiating out from one point. Plant scientists use sophisticated molecular techniques to study the genetic basis of various aspects of plant growth and development, including leaf form. Many of the genes under study code for transcription factors. As discussed in Chapter 14, transcription factors are DNA-binding proteins that regulate the synthesis of 1 Recall that flowering plants, the focus of this chapter, are divided into two main groups, informally called eudicots and monocots (see Chapter 28). Examples of eudicots include beans, petunias, oaks, cherry trees, roses, and snapdragons; monocots include corn, lilies, grasses, palms, tulips, orchids, and bananas.

Simple

Pinnately compound

Palmately compound

White ash (Fraxinus americana)

California white oak (Quercus lobata)

Ohio buckeye (Aesculus glabra)

(a) Leaf form: simple and compound. Alternate

Opposite

Whorled

Sugar maple (Acer saccharum)

American beech (Fagus grandifolia)

Southern catalpa (Catalpa bignonioides )

(b) Leaf arrangement on a stem.

Parallel

Bermuda grass (Cynodon dactylon)

Pinnately netted

Black willow (Salix nigra)

Palmately netted

Sweetgum (Liquidambar styraciflua)

(c) Venation patterns.

FIGURE 34-2 Leaf morphology All leaves shown are woody eudicot trees from North America, except Bermuda grass, which is a herbaceous monocot native to Europe and Asia.

RNA from a DNA template. An exciting example of this type of research was recently published in the journal Science (FIG. 34-3). Eventually, research like this will help plant biologists to understand the roles of all genes involved in plant development—an important goal of systems biology.

Leaf structure is adapted for maximum light absorption The leaf is a complex organ composed of several tissues organized to optimize photosynthesis (FIG. 34-4). The leaf blade has upper

and lower surfaces, each consisting of a layer of epidermis. The upper epidermis covers the upper surface, and the lower epidermis covers the lower surface. Most cells in these layers lack chloroplasts and are relatively transparent. One interesting feature of leaf epidermal cells is that the cell wall facing toward the outside environment is somewhat thicker than the cell wall facing inward. This extra thickness may provide the plant with additional protection against injury or water loss. Because leaves have such a large surface area exposed to the atmosphere, water loss by evaporation from the leaf ’s surface is unavoidable. However, epidermal cells secrete a waxy layer, the cuticle,

KEY EXPERIMENT QUESTION: What molecular controls determine if a developing leaf will become simple or compound? EXPERIMENT: NAM/CUC (NO APICAL MERISTEM/CUP-SHAPED COTYLEDON) genes were cloned and genetically manipulated in four distantly related flowering plants that normally produce compound leaves: Aquilegia caerulea (columbine), Solanum lycopersicum (tomato), Cardamine hirsuta (hairy bittercress), and Pisum sativum (garden pea). NAM/CUC genes were already known to code for plant transcription factors involved in the establishment and function of the shoot apical meristem.

RESULTS AND CONCLUSION: European scientists Thomas Blein, Patrick Laufs, and colleagues reported in the journal Science in 2008 that the timing of gene silencing of NAM/CUC genes had the potential to produce less elaborate, simpler leaf structures. If a young leaf at the shoot apical meristem of a columbine plant had already developed deeply lobed leaflets, silencing the NAM/CUC genes would not affect its further development (leaf 1). The leaf margins of leaves 2 and 3 developed with fewer lobes because the NAM/CUC genes had been silenced before much leaf development had taken place. In leaf 4, very early silencing resulted in a simple leaf with an entire margin. Similar leaf simplifications occurred due to the timing of silencing of the NAM/CUC genes in the developing leaves of tomato, hairy bittercress, and garden pea. Source: Blein, T. et al. “A conserved molecular framework for compound leaf development.” Science, Vol. 322, Dec. 19, 2008. (Photo is taken from Figure 2 on page 1837.)

1

2

3

4

FIGURE 34-3 Molecular controls and the development of leaf form

KEY POINT

Leaves contain all three tissue systems found in plants: the dermal tissue system, represented by the upper and lower epidermis; the ground tissue system, represented by the mesophyll; and the vascular tissue system, represented by the xylem and phloem in the veins.

Palisade mesophyll Vein (vascular bundle)

Cuticle Upper epidermis

Spongy mesophyll

Bundle sheath Xylem Phloem Stoma

Air space Lower epidermis Stoma

FIGURE 34-4 Animated Tissues in a typical leaf blade An upper epidermis and lower epidermis cover the blade. The photosynthetic ground tissue, called mesophyll, is often arranged into palisade and spongy layers. Veins branch throughout the mesophyll.

Guard cells

dermal cells. Subsidiary cells provide a reservoir of water and ions that move into and out of the guard cells as they change shape during stomatal opening and closing (discussed later in the chapter). The photosynthetic ground tissue of the leaf, called the mesophyll (from the Greek meso, “the middle of ”; and phyll, “leaf ”), is sandwiched between the upper epidermis and the lower epidermis. Mesophyll cells, which are parenchyma cells (see Chapter 33) packed with chloroplasts, are loosely arranged, with many air spaces between them that facilitate gas exchange. These intercellular air spaces account for as much as 70% of the leaf ’s volume. In many plants, the mesophyll is divided into two sublayers. Toward the upper epidermis, the columnar cells are stacked closely together in a layer called palisade mesophyll. In the lower portion, the cells are more loosely and irregularly arranged in a layer called spongy mesophyll. The two layers have different functions. Palisade mesophyll is specialized for light capture and is the main site of photosynthesis in the leaf. Photosynthesis also occurs in the spongy mesophyll, but the primary function of the spongy mesophyll is to allow diffusion of gases, particularly CO2, within the leaf. Palisade mesophyll may be organized into one, two, three, or even more layers of cells. The presence of additional layers is at least partly an adaptation to environmental conditions. Leaves exposed to direct sunlight contain more layers of palisade mesophyll than do shaded leaves on the same plant. In direct sunlight, the light is strong enough to effectively penetrate multiple layers of palisade mesophyll, allowing all layers to photosynthesize efficiently. The veins, or vascular bundles, of a leaf extend through the mesophyll. Branching is extensive, and no mesophyll cell is more than two or three cells away from a vein. Therefore, the slow process of diffusion does not limit the movement of needed resources between mesophyll cells and veins. Each vein contains two types of vascular tissue: xylem and phloem. Xylem, which conducts water and dissolved minerals (inorganic nutrients), is usually located in the upper part of a vein, toward the upper epidermis. Phloem, which conducts dissolved sugars, is usually confined to the lower part of a vein. One or more layers of nonvascular cells surround the larger veins and make up the bundle sheath. Bundle sheaths are composed of parenchyma or sclerenchyma cells (see Chapter 33). Frequently, the bundle sheath has support columns, called bundle sheath extensions, that extend through the mesophyll from the upper epidermis to the lower epidermis (FIG. 34-6). Bundle sheath extensions may be composed of parenchyma, collenchyma, or sclerenchyma cells.

that reduces water loss from their exterior walls (see Table 33-4). The cuticle, which consists primarily of the waxy substance cutin, varies in thickness in different plants, in part as a result of environmental conditions. As one might expect, the leaves of plants adapted to hot, dry climates have extremely thick cuticles. Furthermore, a leaf ’s exposed (and warmer) upper epidermis generally has a thicker cuticle than its shaded (and cooler) lower epidermis. The epidermis of many leaves is covered with various hairlike structures called trichomes, which have several functions (see Table 33-4). Trichomes of some plants help reduce water loss from the leaf surface by retaining a layer of moist air next to the leaf and by reflecting sunlight, thereby protecting the plant from overheating. Some trichomes secrete stinging irritants for deterring herbivores, animals that feed on plants. In addition, a leaf covered with trichomes is difficult for an insect to walk over or eat. Other trichomes excrete excess salts absorbed from a salty soil. The leaf epidermis contains minute openings, or stomata (sing., stoma), for gas exchange between leaf cells and the environment; the stomata are evenly spaced to optimize this gas exchange. Each stoma is flanked by two specialized epidermal guard cells, which are responsible for opening and closing the stoma (FIG. 34-5). Guard cells are usually the only epidermal cells with chloroplasts. Stomata are especially numerous on the lower epidermis of horizontally oriented leaves (an average of about 100 stomata per square millimeter) and in many species are located only on the lower surface. The lower epidermis of apple (Malus sylvestris) leaves, for example, has almost 400 stomata per square millimeter, whereas the upper epidermis has none. This adaptation reduces water loss because stomata on the lower epidermis are shielded from direct sunlight and are therefore cooler than those on the upper epidermis. In contrast, floating leaves of aquatic plants, such as water lilies, have stomata only on the upper epidermis. Guard cells are associated with special epidermal cells called subsidiary cells that are often structurally different from other epi-

Subsidiary cells

Guard cells

Epidermal cell Dwight R. Kuhn

Stoma

25 μm

FIGURE 34-5 Stoma LM of an open stoma from the leaf epidermis of an inch plant (Zebrina pendula). Note the chloroplasts present in the guard cells and the thicker inner wall of each guard cell.

Leaf structure differs in eudicots and monocots Each eudicot leaf is usually composed of a broad, flattened blade and a petiole. As mentioned previously, eudicot leaves typically have netted venation. In contrast, many monocot leaves lack a petiole; they are narrow, and the base of the leaf often wraps around the stem to form a sheath. Parallel venation is characteristic of monocot leaves. Eudicots and certain monocots also differ in internal leaf anatomy (FIG. 34-7). Although most eudicots and monocots have both palisade and spongy layers, some monocots (corn and other grasses) do not have mesophyll differentiated into distinct palisade and spongy layers. Because eudicots have netted veins, a cross sec-

▲

FIGURE 34-6 Bundle sheath extension

LM of a wheat (Triticum aestivum) midvein in cross section. Note the bundle sheath extensions to both the upper epidermis and lower epidermis. Photographed using fluorescence microscopy.

Upper epidermis Bundle sheath extension

Midvein Phil Gates/Biological Photo Service

Bundle sheath

Bundle sheath extension Lower epidermis 250 μm

tion of a eudicot blade often shows veins in both cross-sectional and lengthwise views. In a cross section of a monocot leaf, in contrast, the parallel venation pattern produces evenly spaced veins, all of which appear in cross section. Differences between the guard cells in eudicot and certain monocot leaves also occur (FIG. 34-8). The guard cells of eudicots and many monocots are shaped like kidney beans. Other monocot leaves (those of grasses, reeds, and sedges) have guard cells shaped like dumbbells. These structural differences affect how the cells swell or shrink to open or close the stoma.

Upper epidermis

Midvein

Palisade mesophyll Lengthwise view of vein

Spongy mesophyll Ed Reschke/Peter Arnold, Inc.

Air space Lower epidermis

Xylem

Phloem 250 μm

Leaf structure is related to function How is leaf structure related to its primary function of photosynthesis? The epidermis of a leaf is relatively transparent and allows light to penetrate to the interior of the leaf where the photosynthetic ground tissue, the mesophyll, is located. Stomata, which dot the leaf surfaces, permit the exchange of gases between the atmosphere and the leaf ’s internal tissues. CO2, a raw material of photosynthesis, diffuses into the leaf through stomata, and the oxygen produced during photosynthesis diffuses rapidly out of the leaf through stomata. Other gases, including air pollutants, also enter the leaf ’s interior via stomata. Water required for photosynthesis is obtained from the soil and transported in the xylem to the leaf, where it diffuses into the mesophyll and moistens the surfaces of mesophyll cells. The loose arrangement of the mesophyll cells, with air spaces

(a) Eudicot leaf. LM of part of a leaf cross section of privet (Ligustrum vulgare ). The mesophyll has distinct palisade and spongy sections. Midvein Upper epidermis

Mesophyll Bundle sheath cell Air space

Stoma Xylem

Phloem 25 μm

(b) Monocot leaf. LM of part of a leaf cross section of corn (Zea mays). Corn leaves lack distinct regions of palisade and spongy mesophyll.

▲

Dwight R. Kuhn

Lower epidermis

FIGURE 34-7 Eudicot and monocot leaf cross sections

Open

Closed

Pore

Subsidiary cells

Guard cells

(a) Guard cells of eudicots and many monocots are beanshaped. Open

Closed

Pore

Conifers are an important group of woody trees and shrubs that includes pine, spruce, fir, redwood, and cedar.2 Most conifers are evergreen, which means they lose leaves throughout the year rather than during certain seasons. Conifers dominate a large portion of Earth’s land area, particularly in northern forests and mountains. The leaves of most conifers are waxy needles. Their needles have structural adaptations that help them survive winter, the driest part of the year. (Winter is arid even in areas of heavy snows because roots cannot absorb water from soil when the soil temperature is very low.) Indeed, many of the structural features of needles are also found in many desert plants. FIGURE 34-9 shows a cross section of a pine needle. Note that the needle is somewhat thickened rather than thin and bladelike. The needle’s relative thickness, which results in less surface area exposed to the air, reduces water loss. Other features that help conserve water include the thick, waxy cuticle and sunken stomata; these permit gas exchange while minimizing water loss. Thus, needles help conifers tolerate the dry winds that occur during winter (dry refers here to low relative humidity). With the warming of spring, soil water again becomes available, and the needles quickly resume photosynthesis.

Review Guard cells

Subsidiary cells

(b) Some monocot guard cells are narrow in the center and thicker at each end.

■ ■

■

FIGURE 34-8 Variation in guard cells Guard cells are associated with epidermal cells called subsidiary cells.

between cells, allows for rapid diffusion of CO2 to the mesophyll cell surfaces; there it dissolves in a film of water before diffusing into the cells. The veins not only supply the photosynthetic ground tissue with water and minerals (from the roots, by way of the xylem) but also carry (in the phloem) dissolved sugar produced during photosynthesis to all parts of the plant. Bundle sheaths and bundle sheath extensions associated with the veins provide additional support to prevent the leaf, which is structurally weak because of the large amount of air space in the mesophyll, from collapsing under its own weight. Leaves are adapted to help a plant survive in its environment Leaf structure reflects the environment to which a particular plant is adapted. Although both aquatic plants and those adapted to dry conditions perform photosynthesis and have the same basic leaf anatomy, their leaves are modified to enable them to survive different environmental conditions. The leaves of water lilies have petioles long enough to allow the blade to float on the water’s surface. Large air spaces in the mesophyll make the floating blade buoyant. The petioles and other submerged parts have an internal system of air ducts; oxygen moves through these ducts from the floating leaves to the underwater roots and stems, which live in a poorly aerated environment.

■

How are leaves adapted to conserve water? What is the photosynthetic ground tissue of a leaf called? What are its two sublayers? What are the two types of vascular tissue in a vascular bundle? Which vascular tissue is usually located in the upper part of the vascular bundle? How is the leaf organized to deliver the raw materials and remove the products of photosynthesis?

34.2 STOMATAL OPENING AND CLOSING ■ ■ LEARNING OBJECTIVES 5 Explain the role of blue light in the opening of stomata. 6 Outline the physiological changes that accompany stomatal opening and closing.

Stomata are adjustable pores that are usually open during the day when CO2 is required for photosynthesis and closed at night when photosynthesis is shut down (see the section on CAM photosynthesis in Chapter 9 for an interesting exception). The opening and closing of stomata are controlled by changes in the shape of the two guard cells that surround each pore. The guard cells’ shape is determined by their rigidity. When water moves into guard cells from surrounding nonguard cells, the guard cells become turgid (swollen) and bend, producing a pore. When water leaves the guard cells, they become flaccid (limp) and collapse against one 2 As discussed in Chapter 28, conifers are gymnosperms, one of the two groups of seed plants (the other group is the flowering plants, or angiosperms). Unlike flowering plants, whose seeds are enclosed in fruits, conifers bear “naked” seeds on the scales of female cones.

Guard cells of sunken stoma

Epidermis and cuticle

Resin duct Endodermis

John D. Cunningham/ Visuals Unlimited

Xylem Phloem

Vascular bundle

Mesophyll cell (photosynthetic parenchyma cell)

250 μm

FIGURE 34-9 LM of a pine needle in cross section The thick, waxy cuticle and sunken stomata are two structural adaptations that enable pine (Pinus) to retain its needles throughout the winter.

another, closing the pore. What causes water to move into and out of guard cells?

Blue light triggers stomatal opening Data from numerous experiments and observations are beginning to explain the complexities of stomatal movements. Let us begin with stomatal opening, which occurs when the plant detects light from the rising sun. You already know that light is a form of energy; plants absorb light and convert it to chemical energy in the process of photosynthesis. However, light is also an important environmental signal for plants—that is, light provides plants with information about their environment that they use to modify various activities at the molecular and cellular levels. In stomatal opening and several other plant responses, blue light, which has wavelengths from 400 to 500 nm, is an environmental signal. Any plant response to light must involve a pigment, a molecule that absorbs the light, before the induction of a particular biological response. Data such as the responses of stomata to different colors of light suggest that the pigment involved in stomatal opening and closing is yellow (yellow pigments strongly absorb blue light). The yellow pigment is thought to be located in the guard cells, probably in their plasma membranes. In FIGURE 34-10, blue light, which is a component of sunlight, triggers the activation of proton pumps, located in the guard cell plasma membrane. Blue light also triggers the synthesis of malic acid and the hydrolysis (splitting) of starch (discussed later). The proton pumps use ATP energy to actively transport protons (H+) out of the guard cells. The H+ that are pumped are formed when malic acid produced in the guard cells ionizes to form H+ and negatively charged malate ions. As the proton pumps in the plasma membranes of guard cells transport protons out of the guard cells, an electrochemical gradient—that is, a charge

and concentration difference—forms on the two sides of the guard cell plasma membrane. The resulting electrochemical gradient of H+ drives the facilitated diffusion of large numbers of potassium ions into guard cells. This movement occurs through voltage-activated potassium channels, which open when a certain voltage (difference in charge between the two sides of the guard cell plasma membrane) is attained. This movement of potassium ions has been experimentally measured by the patch clamp technique, in which researchers seal the tip of a micropipette to a tiny patch of membrane that contains a single ion channel. They measure the flow of ions through that channel between the cytoplasm and the solution in the micropipette. (Interestingly, in animals, voltage-activated ion channels are found in the plasma membranes of nerve cells and are involved in transmitting neural impulses.) Chloride ions are also taken into the guard cells through ion channels in the guard cell plasma membrane. The negatively charged chloride and malate ions help to electrically balance the positively charged potassium ions. The potassium, chloride, and malate ions accumulate in the vacuoles of the guard cells, thus increasing the solute concentration in the vacuoles. You may recall from the discussion of osmosis in Chapter 5 that when a cell has a solute concentration greater than that of surrounding cells, water flows into the cell. Thus, water enters the guard cells from surrounding epidermal cells by osmosis. The increased turgidity of the guard cells changes their shape, because the thickened inner cell walls do not expand as much as the outer walls, so the stoma opens. blue light activates proton pumps ¡ proton pump moves H+ out of guard cells ¡ K+ and Cl− diffuse into guard cells through voltage-activated ion channels ¡ water diffuses by osmosis into guard cells ¡ guard cells change shape and stoma opens

KEY POINT

The accumulation of osmotically active ions (K+ and Cl−) in the guard cells is driven by a proton (H+) gradient.

H+

1

Blue light activates proton pumps.

2

H+

Protons are pumped out of guard cells, forming electrochemical gradient.

K+

3

Cl-

K+

Potassium ions enter guard cells through voltage-activated ion channels.

4

Cl-

Chloride ions also enter guard cells through ion channels.

H2O

5

H2O

Water enters guard cells by osmosis, and stoma opens.

FIGURE 34-10 Mechanism of stomatal opening

Stomata close in the late afternoon or early evening, but not by an exact reversal of the opening process. Recent studies have demonstrated that the concentration of potassium ions in guard cells slowly decreases during the day. However, the concentration of sucrose, another osmotically active substance, increases during the day—maintaining the open pore—and then slowly decreases as evening approaches. This sucrose comes from the hydrolysis of the polysaccharide starch, which is stored in the guard cell chloroplasts. As evening approaches, the sucrose concentration in the guard cells declines as sucrose is converted back to starch (which is osmotically inactive), water leaves by osmosis, the guard cells lose their turgidity, and the pore closes. To summarize, different mechanisms appear to regulate the opening and closing of stomata. The uptake of potassium and chloride ions is mainly associated with stomatal opening, and the declining concentration of sucrose is mainly associated with stomatal closing. In Chapter 38, we discuss other blue-light responses in plants.

environmental factor that affects stomatal opening and closing is water stress. During a prolonged drought, stomata remain closed, even during the day. Stomatal opening and closing are under hormonal control, particularly by the plant hormone abscisic acid (see Table 38-1). The opening and closing of stomata are also regulated by an internal biological clock that in some way measures time. For example, after plants are placed in continual darkness, their stomata continue to open and close at more or less the same time each day. Such biological rhythms that follow an approximate 24-hour cycle are known as circadian rhythms. Other examples of circadian rhythms are provided in Chapter 38.

Review ■ ■

How does blue light trigger stomatal opening? What physiological changes occur in guard cells during stomatal opening? during stomatal closing?

Additional factors affect stomatal opening and closing

34.3 TRANSPIRATION AND GUTTATION

Although light and darkness trigger the opening and closing of stomata, other environmental factors are also involved, including CO2 concentration. A low concentration of CO2 in the leaf induces stomata to open, even in the dark. The effects of light and CO2 concentration on stomatal opening are interrelated. Photosynthesis, which occurs in the presence of light, reduces the internal concentration of CO2 in the leaf, triggering stomatal opening. Another

■ ■ LEARNING OBJECTIVES 7 8

Discuss transpiration and its effects on plants. Distinguish between transpiration and guttation.

Despite leaf adaptations such as the cuticle, approximately 99% of the water that a plant absorbs from the soil is lost by evaporation

Carlyn Iverson

Carlyn Iverson

from the leaves and, to a lesser extent, the stems. Loss of water vathe soil may not contain sufficient moisture to permit recovery por by evaporation from aerial plant parts is called transpiration. from wilting. A plant that cannot recover is said to be permanently The cuticle is extremely effective in reducing water loss by tranwilted and will die. spiration. It is estimated that only 1% to 3% of the water lost from a Transpiration is an important part of the hydrologic cycle, in plant passes directly through the cuticle. Most transpiration occurs which water cycles from the ocean and land to the atmosphere and through open stomata. The numerous stomatal pores that are so then back to the ocean and land (see Fig. 55-11). As a result of traneffective in gas exchange for photosynthesis also provide openings spiration, water evaporates from leaves and stems to form clouds in through which water vapor escapes. In addition, the loose arrangethe atmosphere. Thus, transpiration eventually results in precipitament of the spongy mesophyll cells provides a large surface area tion. As you may expect, forest trees release substantial amounts of within the leaf from which water can evaporate. moisture into the air by transpiration. Researchers have determined Several environmental factors influence the rate of transpirathat at least half the rain that falls in the Amazon rainforest basin is tion. More water is lost from plant surfaces at higher temperatures. recycled again and again by transpiration and precipitation. Light increases the transpiration rate, in part because it triggers stomatal opening and in part because it increases the leaf ’s temSome plants exude liquid water perature. Wind and dry air increase transpiration, but humid air decreases transpiration because the air is already saturated, or nearly Many leaves have special structures at vein endings through which so, with water vapor. liquid water is literally forced out. This loss of liquid water, known Although transpiration may seem wasteful, particularly to as guttation, occurs when transpiration is negligible and available farmers in arid lands, it is an essential process that has adaptive soil moisture is high. Guttation typically occurs in low-growing value. Transpiration is responsible for water movement in plants, plants at night because the stomata are closed, but water continues and without it water would not reach the leaves from the soil (see to move into the roots by osmosis. People sometimes mistake the discussion of the tension–cohesion model in Chapter 35). The early morning water droplets formed on leaf margins by guttation large amount of water that plants lose by transpiration may profor dew (FIG. 34-12). Unlike dew, which condenses from cool night vide some additional benefits. Transpiration, like sweating in huair, guttation droplets come from within the plant. (The mechamans, cools the leaves and stems. When water passes from a liquid nism for guttation is discussed in Chapter 35.) state to a vapor, it absorbs a great deal of heat. When the water molecules leave the plant as water vapor, they carry this heat with Review them. Thus, the cooling effect of transpiration may prevent leaves from overheating, particularly in direct sunlight. ■ What is transpiration? How is leaf structure related to transpiration? Another benefit of transpiration is that it distributes essential ■ How do environmental factors (sunlight, temperature, humidity, and wind) influence the rate of transpiration? minerals throughout the plant. The water that a plant transpires ■ How does guttation differ from transpiration? is initially absorbed from the soil, where it is present not as pure water but as a dilute solution of dissolved mineral salts. The water and dissolved minerals are then transported in the xylem throughout the plant, including its leaves. Water moves from the plant to the atmosphere during transpiration, but minerals remain in plant tissues. Many of these minerals are required for the plant’s growth. It has been suggested that transpiration enables a plant to take in sufficient water to provide enough essential minerals; it has also been suggested that plants cannot satisfy their mineral requirements if the transpiration rate is not high enough. There is no doubt, however, that under certain circumstances excessive transpiration can be harmful to a plant. On hot summer days, plants frequently lose more water by transpiration than they can take in from the soil. Their cells experience a loss of turgor, and the plants wilt (FIG. 34-11). If a plant is able to recover over(b) The next morning, water in the leaves (a) In the late afternoon of a hot day, night, because of the combination of negligible has been replenished. Transpiration is the leaves have wilted because of water transpiration (recall that stomata are closed) and negligible during the night, and the loss. Note that wilting helps reduce the absorption of water from the soil, the plant is plants recover by absorbing water from surface area from which transpiration the soil. occurs. said to have experienced temporary wilting. Most plants recover from temporary wilting with no ill FIGURE 34-11 Temporary wilting in squash (Cucurbita pepo) leaves effects. In cases of prolonged drought, however,

are mobilized and transported from the leaves to other plant parts; these minerals will be reused during the following spring. Chlorophyll breaks down, allowing the carotenoids (orange carotenes and yellow xanthophylls), some of the accessory pigments in the chloroplasts of leaf cells, to become evident (see Fig. 3-14b). Accessory pigments are always present in the chloroplasts but are masked by the green of the chlorophyll. In addition, red water-soluble pigments called anthocyanins may accumulate in the vacuoles of epidermal leaf cells in some species; anthocyanins may protect leaves against damage by ultraviolet radiation. The various combinations of carotenoids and anthocyanins are responsible for the brilliant colors found in autumn landscapes in temperate climates.

Ed Reschke/Peter Arnold, Inc.

In many leaves, abscission occurs at an abscission zone near the base of the petiole

FIGURE 34-12 Guttation Shown is a compound leaf of strawberry (Fragaria) with water droplets formed by guttation.

34.4 LEAF ABSCISSION ■ ■ LEARNING OBJECTIVE 9 Define leaf abscission, explain why it occurs, and describe the physiological and anatomical changes that precede it.

All trees shed leaves, a process known as abscission. Many conifers shed their needles in small numbers year-round. The leaves of deciduous plants turn color and abscise, or fall off, once a year— as winter approaches in temperate climates or at the beginning of the dry period in tropical climates with pronounced wet and dry seasons. In temperate forests, most woody plants with broad leaves shed their leaves to survive the low temperatures of winter. During winter, the plant’s metabolism, including its photosynthetic machinery, slows down or halts temporarily. Another reason for abscission is related to a plant’s water requirements, which become critical during the physiological drought of winter. As mentioned previously, as the ground chills, absorption of water by the roots is inhibited. When the ground freezes, no absorption occurs. If the broad leaves were to stay on the plant during the winter, the plant would continue to lose water by transpiration but would be unable to replace it with water absorbed from the soil. Leaf abscission is a complex process that involves many physiological changes, all initiated and orchestrated by changing levels of plant hormones, particularly ethylene. Briefly, the process is this: As autumn approaches, sugars, amino acids, and many essential minerals (such as nitrogen, phosphorus, and possibly potassium)

The area where a petiole detaches from the stem is a structurally distinct area called the abscission zone. Composed primarily of thin-walled parenchyma cells, it is anatomically weak because it contains few fibers (FIG. 34-13). A protective layer of cork cells develops on the stem side of the abscission zone. These cells have a waxy, waterproof material impregnated in their walls. Enzymes then dissolve the middle lamella (the “cement” that holds the primary cell walls of adjacent cells together) in the abscission zone (see Fig. 4-29). Once this process is completed, nothing holds the leaf to the stem but a few xylem cells. A sudden breeze is enough to make the final break, and the leaf detaches. The protective layer of cork remains, sealing off the area and forming a leaf scar. Although the structure of abscission zones and the physiological changes associated with abscission have been well known for many years, only recently have botanists investigated the genes responsible for the development of an abscission zone and the physiological changes of leaf abscission. Using DNA microarrays, for example, botanists have identified about 35 genes that are expressed in leaves only as winter approaches. As shown in Figure 15-12, a DNA microarray contains hundreds of different DNA molecules placed on a glass slide or chip. Some of these genes code for degradative enzymes involved in breaking down proteins and other molecules; others have functions not yet identified.

Review ■

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Why do many woody plants living in temperate zones lose their leaves in autumn? What physiological changes occur during leaf abscission? What is the abscission zone?

34.5 MODIFIED LEAVES ■ ■ LEARNING OBJECTIVE 10 List at least four examples of modified leaves and give the function of each.

Although photosynthesis is the main function of leaves, certain leaves have special modifications for other functions. Some plants have leaves specialized for deterring herbivores. Spines, modified

Axillary bud

Bud scales

FIGURE 34-13 Abscission zone This LM of a longitudinal section through a silver maple (Acer saccharinum) branch shows the abscission zone at the base of a petiole. An axillary bud with its protective bud scales is also evident above the petiole.

James Mauseth, University of Texas

Petiole

leaves that are hard and pointed, are found on many desert plants, such as cacti (FIG. 34-14a). In the cactus, the main organ of photosynthesis is the stem rather than the leaf. Spines discourage animals from eating the succulent stem tissue. Vines are climbing plants whose stems cannot support their own weight, so they often possess tendrils that help keep the vine attached to the structure on which it is growing. The tendrils of many vines, such as peas, cucumbers, squash, and sweet peas, are specialized leaves (FIG. 34-14b). However, some tendrils, such as those of ivy, Virginia creeper, and grape, are specialized stems. The winter buds of a dormant woody plant are covered by bud scales, modified leaves that protect the delicate meristematic tissue of the bud from injury, freezing, or drying out (FIG. 34-14c). Some leaves associated with flower clusters (inflorescences) are modified as bracts. In flowering dogwood, the inconspicuous flowers are clustered in the center of each inflorescence, and what appears to be four white or pink petals are actually bracts. Similarly, the red “petals” of poinsettia are not petals at all but bracts (FIG. 34-14d). Leaves may also be modified for storage of water or food. For example, a bulb is a short, underground stem to which large, fleshy leaves are attached (FIG. 34-14e). Onions and tulips form bulbs. Many plants adapted to arid conditions, such as jade plant, medicinal aloe, and string-of-beads, have succulent leaves for water storage (FIG. 34-14f). These leaves are usually green and function in photosynthesis. Some leaves are modified for either sexual reproduction (see discussion of flower evolution in Chapter 28, including Fig. 28-15) or asexual reproduction (see Fig. 37-17).

Abscission zone

Stem 0.5 mm

Modified leaves of carnivorous plants capture insects Carnivorous plants are plants that capture insects. Most carnivorous plants grow in poor soil that is deficient in certain essential minerals, particularly nitrogen. These plants meet some of their mineral requirements by digesting insects and other small animals. The leaves of carnivorous plants are adapted to attract, capture, and digest their animal prey. Some carnivorous plants have passive traps. The leaves of a pitcher plant, for example, are shaped so that rainwater collects and forms a reservoir that also contains acid secreted by the plant (FIG. 34-15). Some pitchers are quite large; in the tropics, pitcher plants may be large enough to hold 1 L (approximately 1 qt) or more of liquid. An insect attracted by the odor or nectar of the pitcher may lean over the edge and fall in. Although it may make repeated attempts to escape, the insect is prevented from crawling out by the slippery sides and the rows of stiff hairs that point downward around the lip of the pitcher. The insect eventually drowns, and part of its body disintegrates and is absorbed. Most insects are killed in pitcher plants. However, the larvae of several insects (certain flies, midges, and mosquitoes) and a large community of microorganisms live inside the pitchers. These insect species obtain their food from the insect carcasses, and the pitcher plant digests what remains. It is not known how these insects survive the acidic environment inside the pitcher. The Venus flytrap is a carnivorous plant with active traps. Its leaf blades resemble tiny bear traps (see Fig. 1-3). Each side of the leaf blade contains three small, stiff hairs. If an insect alights and

Cengage

© Mark Bolton/ Terra/Corbis

Barbara Gerlach / Dembinsky Photo Associates

(b) Tendrils of a sweet pea (Lathyrus odoratus) are modified leaves that wind around objects and aid in climbing.

(c) A terminal bud and two axillary buds of a maple (Acer) twig have overlapping bud scales to protect the buds.

Cengage

© Radius Images/Corbis

(a) The leaves of this pincushion cactus (Mammilaria) are spines for deterring herbivores.

(e) The leaves of bulbs such as the onion (Allium cepa ) are fleshy for storage of food materials and water.

James Mauseth, University of Texas

(d) Showy red bracts surround each poinsettia (Euphorbia pulcherrima) inflorescence; the inconspicuous flowers are in the center.

(f) The succulent leaves of the string-of-beads plant (Senecio rowleyanus) are spherical to minimize surface area, thereby conserving water.

FIGURE 34-14 Leaf modifications

▲

FIGURE 34-15 A common pitcher plant

© Marvin Dembinsky Photo Associates/Alamy

This species (Sarracenia purpurea), whose pitchers grow to 30.5 cm (12 in), is widely distributed in acidic bogs and marshes in eastern North America. Young pitchers are green but turn red as they age. Note the dead beetle in the pitcher.

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brushes against two of the hairs, or against the same hair twice in quick succession, the trap springs shut with amazing rapidity— about 100 milliseconds. The spines along the margins of the blades fit closely together to prevent the insect from escaping. After the leaf initially traps the insect, the leaf continues to slowly close for the next several hours. Digestive glands on the surface of the trap secrete enzymes in response to the insect pressing against them. Days later, after the insect has died and been digested, the trap reopens and the indigestible remains fall out.

Review What are the primary functions of each of the following modified leaves: spines, tendrils, and bud scales? What are the functions of bulbs? of succulent leaves?

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What are some of the specialized features of the leaves of carnivorous plants?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

34.1 (page 729)

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1 Discuss variation in leaf form, including simple versus compound leaves, leaf arrangement on the stem, and venation patterns. Leaves may be simple (having a single blade) or compound (having a blade divided into two or more leaflets). ■ Leaf arrangement on a stem may be alternate (one leaf at each node), opposite (two leaves at each node), or whorled (three or more leaves at each node). ■ Leaves may have parallel or netted venation. Netted venation may be palmately netted, with several major veins radiating from one point, or pinnately netted, with veins branching along the entire length of the midvein. 2 Describe the major tissues of the leaf (epidermis, photosynthetic ground tissue, xylem, and phloem) and label them on a diagram of a leaf cross section. ■ Upper and lower surfaces of the leaf blade are covered by an epidermis. A waxy cuticle coats the epidermis, enabling the plant to survive the dry conditions of a terrestrial existence. ■ Stomata are small pores in the epidermis that permit gas exchange needed for photosynthesis. Each pore is surrounded by two guard cells that are often associated with special epidermal cells called subsidiary cells. Subsidiary cells provide a reservoir of water and ions that move into and out of the guard cells as they change shape during stomatal opening and closing. ■ Mesophyll consists of photosynthetic parenchyma cells. Mesophyll is divided into palisade mesophyll, which functions primarily for photosynthesis, and spongy mesophyll, which functions primarily for gas exchange. ■

Leaf veins have xylem to conduct water and essential minerals to the leaf and phloem to conduct sugar produced by photosynthesis to the rest of the plant. Learn more about leaf tissues by clicking on the figure in CengageNOW.

3 Compare leaf anatomy in eudicots and monocots. Monocot leaves have parallel venation, whereas eudicot leaves have netted venation. Some monocots (corn and other grasses) do not have mesophyll differentiated into distinct palisade and spongy layers. Some monocots (grasses, reeds, and sedges) have guard cells shaped like dumbbells, unlike the more common bean-shaped guard cells. 4 Relate leaf structure to its function of photosynthesis. ■ Leaf structure is adapted for its primary function of photosynthesis. Most leaves have a broad, flattened blade that efficiently collects the sun’s radiant energy. The transparent epidermis allows light to penetrate into the middle of the leaf, where photosynthesis occurs. ■ Stomata generally open during the day for gas exchange needed during photosynthesis and close at night to conserve water when photosynthesis is not occurring. ■ Air spaces in mesophyll tissue permit the rapid diffusion of CO2 and water into, and oxygen out of, mesophyll cells. ■

34.2 (page 734) 5 Explain the role of blue light in the opening of stomata. ■

Blue light, which is a component of sunlight, triggers the activation of proton pumps located in the guard cell plasma membrane. Blue

light also triggers the synthesis of malic acid and the hydrolysis of starch.

34.3 (page 736) 7 Discuss transpiration and its effects on plants. Transpiration is the loss of water vapor from aerial parts of plants. Transpiration, which occurs primarily through the stomata, is affected by environmental factors such as temperature, wind, and relative humidity. ■ Transpiration appears to be both beneficial and harmful to the plant—that is, transpiration represents a trade-off between the CO2 requirement for photosynthesis and the need for water conservation. 8 Distinguish between transpiration and guttation. ■ Guttation, the release of liquid water from leaf margins of some plants, occurs through special structures when transpiration is negligible and available soil moisture is high. In contrast, transpiration is the loss of water vapor and occurs primarily through the stomata. ■

34.4 (page 738) 9 Define leaf abscission, explain why it occurs, and describe the physi6 Outline the physiological changes that accompany stomatal opening and closing. Protons (H+), produced when malic acid ionizes, are pumped out of the guard cells. As protons leave the guard cells, an electrochemical gradient (a charge and concentration difference) forms on the two sides of the guard cell plasma membrane. ■ The electrochemical gradient drives the uptake of potassium ions (K+) through voltage-activated potassium channels into the guard cells. Chloride ions (C−) are also taken into the guard cells through ion channels. These osmotically active ions increase the solute concentration in the guard cell vacuoles. The resulting osmotic movement of water into the guard cells causes them to become turgid, forming a pore. ■ As the day progresses, potassium ions slowly leave the guard cells and starch is hydrolyzed to sucrose, an osmotically active solute, which increases in concentration in the guard cells. Stomata close when water leaves the guard cells as a result of a decline in the concentration of sucrose, which occurs as evening approaches. The sucrose is converted to starch, which is osmotically inactive. ■

Watch stomata in action by clicking on the figure in CengageNOW.

ological and anatomical changes that precede it. ■ Leaf abscission is the loss of leaves that often occurs as winter approaches in temperate climates or at the beginning of the dry period in tropical climates with wet and dry seasons. ■ Abscission involves physiological and anatomical changes that occur prior to leaf fall. An abscission zone develops where the petiole detaches from the stem. Sugars, amino acids, and many essential minerals are transported from the leaves to other plant parts. Chlorophyll breaks down, and carotenoids and anthocyanins become evident.

34.5 (page 738) 10 List at least four examples of modified leaves and give the function of each. Spines are leaves adapted to deter herbivores. Some tendrils are leaves modified for grasping and holding on to other structures (to support weak stems). Bud scales are leaves modified to protect delicate meristematic tissue or dormant buds. Bracts are modified leaves associated with some inflorescences. Bulbs are short, underground stems with fleshy leaves specialized for storage. Many plants adapted to arid conditions have succulent leaves for water storage. Carnivorous plants have leaves modified to trap insects.

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T E S T YO U R U N D E R S TA N D I N G 1. Plants with an alternate leaf arrangement have (a) blades divided into two or more leaflets (b) major veins that radiate out from one point (c) one leaf at each node (d) major veins branching off along the entire length of the midvein (e) two leaves at each node 2. The primary function of the spongy mesophyll is (a) reducing water loss from the leaf surface (b) changing the shape of the guard cells (c) supporting the leaf to prevent it from collapsing under its own weight (d) diffusing gases within the leaf (e) deterring herbivores 3. Gas exchange occurs through microscopic pores formed by two (a) subsidiary cells (b) abscission cells (c) mesophyll cells (d) guard cells (e) stipules 4. The thin, noncellular layer of wax secreted by the epidermis of leaves is the (a) stoma (b) subsidiary cell (c) trichome (d) bundle sheath (e) cuticle

5. Which of the following is not an adaptation of pine needles to conserve water? (a) less surface area exposed to the air than thinbladed leaves (b) a relatively thick cuticle (c) sunken stomata (d) netted veins instead of parallel veins (e) both c and d are not adaptations of pine needles 6. When transpiration is negligible, plants such as grasses exude excess water by (a) guttation (b) circadian rhythm (c) abscission (d) pumping H+ out of and K+ into guard cells (e) photosynthesis 7. At sunrise, the accumulation in the guard cells of the osmotically active substance _____ causes an inflow of water and the opening of the pore. (a) protons (b) starch (c) ATP synthase (d) sucrose (e) potassium ions 8. Stomatal opening is most pronounced in response to _____ light. (a) green (b) yellow (c) blue (d) ultraviolet (e) infrared

9. Anatomically, the abscission zone where a petiole detaches from a stem consists of (a) thin-walled parenchyma cells with few fibers (b) thick-walled cork parenchyma cells (c) clusters of fibers and collenchyma strands (d) hard, pointed stipules (e) epidermal cells with sunken stomata 10. Modified leaves that enable a stem to climb are called _____ , whereas modified leaves that cover the winter buds of a dormant woody plant are called _____. (a) spines; bud scales (b) bud scales; tendrils (c) tendrils; bud scales (d) tendrils; spines (e) carnivorous leaves; spines

11. There is a trade-off between photosynthesis and transpiration in leaves because (a) numerous stomatal pores provide both gas exchange for photosynthesis and openings through which water vapor escapes (b) a waxy layer, the cuticle, reduces water loss (c) blue light triggers an influx of potassium ions (K+) into the guard cells (d) leaves of deciduous plants abscise as winter approaches in temperate climates (e) stomata are closed at night, although water continues to move into the roots by osmosis 12. Label the following diagram. (Refer to Figure 34-4 to check your answers.)

CRITICAL THINKING 1. Suppose that you are asked to observe a micrograph of a leaf cross section and distinguish between the upper and lower epidermis. How would you make this decision? 2. Given that (a) xylem is located toward the upper epidermis in leaf veins and phloem is toward the lower epidermis and (b) the vascular tissue of a leaf is continuous with that of the stem, suggest one possible arrangement of vascular tissues in the stem that might account for the arrangement of vascular tissue in the leaf. 3. What might be some of the advantages of a plant having a few large leaves? What might be some disadvantages? What might be some advantages of having many small leaves? What disadvantages might this entail? How would your answers differ for plants growing in a humid environment compared to those in a desert?

4. EVOLUTION LINK. Why did natural selection favor the evolution of seasonal leaf abscission in woody flowering plants living in colder climates? What adaptations enable conifers to survive these climates without leaf abscission? 5. SCIENCE, TECHNOLOGY, AND SOCIETY. Briefly explain why research on the molecular mechanism of stomatal closure might be of future use in agriculture. Additional questions are available in CengageNOW at www.cengage.com/ login.

35

Stem Structure and Transport

Gregory G. Dimijian, M.D./Photo Researchers, Inc.

Giant sequoias in winter. Giant sequoias (Sequoiadendron giganteum) are the most massive trees on Earth. Their huge stems grow to 75 m (250 ft) or more. Photographed in Sequoia National Park, California.

KEY CONCEPTS

A

35.1 Primary tissues (epidermis, cortex, pith, xylem, and

anchor the plant and absorb materials from the soil, whereas leaves are

phloem) of stems develop from shoot apical meristems. Secondary tissues (wood and bark) of stems develop from two lateral meristems, vascular cambium and cork cambium.

primarily for photosynthesis, converting radiant energy into the chemical

35.2 The concept of water potential explains the direction of water transport into, through, and out of a plant. According to the tension–cohesion model, transpiration pulls water up through the stem as water evaporates from leaves by transpiration.

35.3 According to the pressure–flow model, sucrose is translocated in phloem sap from the source, where the sugar is loaded into phloem, to the sink, where the sugar is removed from phloem.

vegetative (not sexually reproductive) vascular plant has three parts: roots, leaves, and stems. As discussed in Chapter 33, roots serve to

energy of carbohydrate molecules. Stems, the focus of this chapter, link a plant’s roots to its leaves and are usually located aboveground, although many plants have underground stems. Stems exhibit varied forms, ranging from ropelike vines to massive tree trunks (see photograph). They can be either herbaceous, with soft, nonwoody tissues, or they can be woody, with extensive hard tissues of wood and bark. Stems perform three main functions in plants. First, stems of most species support leaves and reproductive structures. The upright position of most stems and the arrangement of the leaves on them allow each leaf to absorb light for use in photosynthesis. Reproductive structures (such

as flowers and fruits) are located on stems in areas accessible to insects, birds, and air currents, which transfer pollen from flower to flower and help disperse seeds and fruits. Second, stems provide internal transport. They conduct water and dissolved minerals (inorganic nutrients) from the roots, where these materials are absorbed from the soil, to leaves and other plant parts. Stems also conduct the sugar produced in leaves by photosynthesis to roots and other parts of the plant. Remember, however, that stems are not the only plant organs that conduct materials. The vascular system is continuous throughout all parts of a plant, and conduction occurs in roots, stems, leaves, and reproductive structures. Third, stems produce new living tissue. They continue to grow throughout a plant’s life, producing buds that develop into stems with new leaves and/or reproductive structures. In addition to the main functions of support, conduction, and production of new stem tissues, stems of some species are modified for asexual reproduction (see Chapter 37) or, if green, to manufacture sugar by photosynthesis.

35.1 STEM GROWTH AND STRUCTURE ■ ■ LEARNING OBJECTIVES 1 2 3

Label cross sections of herbaceous eudicot and monocot stems, and describe the functions of each tissue. Name the two lateral meristems and describe the tissues that arise from each. Outline the transition from primary growth to secondary growth in a woody stem.

You may recall from Chapter 33 that plants have two different types of growth. Primary growth is an increase in the length of a plant and occurs at apical meristems located at the tips of roots and shoots and also within the buds of stems. Secondary growth is an increase in the girth (thickness) of a plant as a result of the activity of lateral meristems located within stems and roots. The new tissues formed by the lateral meristems are called secondary tissues to distinguish them from primary tissues produced by apical meristems. All plants have primary growth; some plants have both primary and secondary growth. Stems with only primary growth are herbaceous, whereas those with both primary and secondary growth are generally woody. (Certain herbaceous stems, such as geranium and sunflower, also have a limited amount of secondary growth.) A woody plant increases in length by primary growth at the tips of its stems and roots, whereas its older stems and roots farther back from the tips increase in girth by secondary growth. In other words, at the same time that primary growth is increasing the length of the stem, secondary growth is adding wood and bark, thereby causing the stem to thicken.

Herbaceous eudicot and monocot stems differ in internal structure Although considerable structural variation exists in stems, they all possess an outer protective covering (epidermis or periderm), one or more types of ground tissue, and vascular tissues (xylem and phloem). Let us first consider the structure of herbaceous eudicot stems and then of monocot stems. Vascular bundles of herbaceous eudicot stems are arranged in a circle in cross section A young sunflower stem is a representative herbaceous eudicot stem that exhibits primary growth (FIG. 35-1). Its outer covering, the epidermis, provides protection in herbaceous stems, as it does in leaves and herbaceous roots (see Table 33-4 and Fig. 33-5). The cuticle, a waxy layer of cutin, covers the stem epidermis and reduces water loss from the stem surface. Stomata permit gas exchange. (Recall from Chapter 34 that a cuticle and stomata are also associated with the leaf epidermis.) Inside the epidermis is the cortex, a cylinder of ground tissue that may contain parenchyma, collenchyma, and sclerenchyma cells (see Table 33-2 and Fig. 33-3). As might be expected from the various types of cells that it contains, the cortex in herbaceous eudicot stems can have several functions, such as photosynthesis, storage, and support. If a stem is green, photosynthesis occurs in chloroplasts of cortical parenchyma cells. Parenchyma in the cortex also stores starch (in amyloplasts) and crystals (in vacuoles). Collenchyma and sclerenchyma in the cortex confer strength and structural support for the stem. The vascular tissues provide conduction and support. In herbaceous eudicot stems, the vascular tissues are located in bundles that, when viewed in cross section, are arranged in a circle. However, viewed lengthwise, these bundles extend as long strands throughout the length of a stem and are continuous with vascular tissues of both roots and leaves. Each vascular bundle contains both xylem, which transports water and dissolved minerals from roots to leaves, and phloem, which transports dissolved sugar (see Table 33-3 and Fig. 33-4). Xylem is located on the inner side of the vascular bundle, and phloem is found toward the outside. Sandwiched between xylem and phloem in some herbaceous stems is a single layer of cells called the vascular cambium, a lateral meristem responsible for secondary growth (discussed later). Because most stems support the aerial plant body, they are much stronger than roots. The thick walls of tracheids and vessel elements in xylem help support the plant. Fibers also occur in both xylem and phloem, although they are usually more extensive in phloem. These fibers add considerable strength to the herbaceous stem. In sunflowers and certain other herbaceous eudicot stems, phloem contains a cluster of fibers toward the outside of the vascular bundle, called a phloem fiber cap, that helps strengthen the stem. The phloem fiber cap is not present in all herbaceous eudicot stems. The pith is a ground tissue at the center of the herbaceous eudicot stem that consists of large, thin-walled parenchyma cells that function primarily in storage. Because of the arrangement of the vascular tissues in bundles, there is no distinct separation of cortex

Epidermis Vascular bundles

Cortex Phloem fiber cap Phloem

Epidermis

Vascular cambium

Ed Reschke/Peter Arnold, Inc.

c. Ed Re schke/ Peter Arnold, In

Cortex

Xylem Vessel element Pith

500 μm

(a) Cross section of a sunflower (Helianthus annuus) stem. Note the vascular bundles arranged in a circle around a central core of pith.

Vascular bundle

Pith

250 μm

(b) Close-up of a vascular bundle. In each bundle, xylem is located toward the stem’s interior, and phloem toward the exterior. Each vascular bundle is “capped” by a batch of fibers for additional support.

FIGURE 35-1 Animated LMs of a herbaceous eudicot stem

and pith between the vascular bundles. The areas of parenchyma between the vascular bundles are often referred to as pith rays.

Woody plants have stems with secondary growth

Vascular bundles are scattered throughout monocot stems An epidermis with its waxy cuticle covers monocot stems, such as the herbaceous stem of corn. As in herbaceous eudicot stems, the vascular tissues run in strands throughout the length of a stem. In cross section the vascular bundles contain xylem toward the inside and phloem toward the outside. In contrast with herbaceous eudicots, however, vascular bundles of monocots are not arranged in a circle but are scattered throughout the stem (FIG. 35-2). Each vascular bundle is enclosed in a bundle sheath of supporting sclerenchyma cells. The monocot stem does not have distinct areas of cortex and pith. The ground tissue in which the vascular tissues are embedded performs the same functions as cortex and pith in herbaceous eudicot stems. Monocot stems do not possess lateral meristems (vascular cambium and cork cambium) that give rise to secondary growth. Monocots have primary growth only and do not produce wood and bark. Although some treelike monocots (such as palms) attain considerable size, they do so by a modified form of primary growth in which parenchyma cells divide and enlarge. Stems of some monocots (such as bamboo and palm) contain a great deal of sclerenchyma tissue, which makes them hard and woodlike in appearance.

Woody plants undergo secondary growth, an increase in the girth of stems and roots. Secondary growth occurs as a result of the activity of two lateral meristems: vascular cambium and cork cambium. Among flowering plants, only woody eudicots (such as apple, hickory, and maple) have secondary growth. Cone-bearing gymnosperms (such as pine, juniper, and spruce) also have secondary growth. Cells in the vascular cambium divide and produce two conducting and supporting tissues: secondary xylem (wood) to replace primary xylem and secondary phloem (inner bark) to replace primary phloem. Primary xylem and primary phloem are not able to transport materials indefinitely and so are replaced in plants that have extended lifespans. Cells of the outer lateral meristem, called cork cambium, divide and produce cork cells and cork parenchyma. Cork cambium and the tissues it produces are collectively referred to as periderm (outer bark), which functions as a replacement for the epidermis (see the last LM in Table 33-4). Vascular cambium gives rise to secondary xylem and secondary phloem Primary tissues in woody eudicot stems are organized like those in herbaceous eudicot stems, with the vascular cambium a thin layer of cells sandwiched between xylem and phloem in the vascular

Phloem

Ground tissue Vascular bundles

Sieve tube element Companion cell Xylem

Epidermis 500 μm

(a) Cross section of a corn (Zea mays) stem shows vascular bundles scattered throughout ground tissue.

Ed Reschke/Peter Arnold, Inc.

Ed Reschke

Vessel element Air space Bundle sheath (surrounds the vascular bundle) 100 μm

(b) Close-up of a vascular bundle. The air space is the site where the first xylem elements were formed and later disintegrated. The entire bundle is enclosed in a bundle sheath of sclerenchyma for additional support.

FIGURE 35-2 Animated LMs of a monocot stem

bundles. Once secondary growth begins, the internal structure of a stem changes considerably (FIG. 35-3). Although vascular cambium is not initially a continuous cylinder of cells (because the vascular bundles are separated by pith rays), it becomes continuous when production of secondary tissues begins. This continuity develops because certain parenchyma cells in each pith ray retain the ability to divide. These cells connect to vascular cambium cells in each vascular bundle, forming a complete ring of vascular cambium. Cells in the vascular cambium divide and produce daughter cells in two directions. The cells formed from the dividing vascular cambium are located either inside the ring of vascular cambium (to become secondary xylem, or wood) or outside it (to become secondary phloem, or inner bark) (FIG. 35-4). When a cell in the vascular cambium divides tangentially (inward or outward), one daughter cell remains meristematic; that is, it remains part of the vascular cambium. The other cell may divide again several times, but eventually it stops dividing and develops into mature secondary tissue. Thus, vascular cambium is a thin layer of cells sandwiched between the wood and inner bark, the two tissues it produces (FIG. 35-5). As the stem increases in circumference, the number of cells in the vascular cambium also increases. This increase in cells occurs by an occasional radial division of a vascular cambium cell, at right angles to its normal direction of division. In this case, both daughter cells remain meristematic.

What happens to the original primary tissues of a stem once secondary growth develops? As a stem increases in thickness, the orientation of the original primary tissues changes. For example, secondary xylem and secondary phloem are laid down between the primary xylem and primary phloem within each vascular bundle. Therefore, as vascular cambium forms secondary tissues, the primary xylem and primary phloem in each vascular bundle become separated from one another. The primary tissues located outside the cylinder of secondary growth (that is, primary phloem, cortex, and epidermis) are subjected to the mechanical pressures produced by secondary growth and are gradually crushed or torn apart and sloughed off. Secondary tissues replace the primary tissues in function. Secondary xylem conducts water and dissolved minerals from roots to leaves in the woody plant. It contains the same types of cells found in primary xylem: water-conducting tracheids and vessel elements, in addition to xylem parenchyma cells and fibers. The arrangement of the different cell types in secondary xylem produces the distinctive wood characteristics of each species. Secondary phloem conducts dissolved sugar from its place of manufacture or of breakdown of starch to a place of use and storage. The same types of cells found in primary phloem (sieve tube elements, companion cells, phloem parenchyma cells, and fibers) are also found in secondary phloem, although secondary phloem usually has more fibers than primary phloem.

KEY POINT

The stems of many plants undergo secondary growth to increase in girth.

Primary xylem

Epidermis Cortex

Primary phloem

Vascular cambium Pith 1

At the onset of secondary growth, vascular cambium arises in the parenchyma between the vascular bundles (that is, in the pith rays), forming a cylinder of meristematic tissue (blue circle in cross section).

Remnant of primary phloem

Remnant of cortex

Remnant of epidermis Secondary phloem (inner bark) Secondary xylem (wood) Periderm (outer bark)

Remnant of primary xylem 2

Remnant of pith

Vascular cambium

Vascular cambium begins to divide, forming secondary xylem on the inside and secondary phloem on the outside.

Secondary xylem (wood)

Periderm (outer bark; remnants of primary phloem, cortex and epidermis are gradually crushed or torn apart and sloughed off) Secondary phloem (inner bark)

Remnant of primary xylem 3

Remnant of pith

Vascular cambium

A young woody stem. Vascular cambium produces significantly more secondary xylem than secondary phloem.

FIGURE 35-3 Development of secondary growth Vascular cambium and the tissues it produces are shown in cross section; cork cambium is not depicted. (The figures change in scale because of space limitations; pith and primary xylem are actually the same size in all three diagrams, but the change in scale makes those tissues appear to shrink in each succeeding diagram.)

Secondary xylem and secondary phloem transport water, minerals, and sugar vertically throughout the woody plant body. However, materials must also move horizontally (laterally). Lateral movement occurs through rays, which are chains of parenchyma cells that radiate from the center of the woody stem or root

(see Fig. 35-5). The vascular cambium forms rays, which are often continuous from the secondary xylem to the secondary phloem. Water and dissolved minerals are transported laterally through rays, from the secondary xylem to the secondary phloem. Likewise, rays form pathways for the lateral transport of dissolved sugar, from the

▲

FIGURE 35-4 Animated Divisions of a vascular cambium cell during a single growing season

X1 X2 X3 X4

P2 P1

X1 X2 Time

P2 P1

P2 P1

X1 X2 X3

Secondary xylem

Secondary phloem P1

X1 X2

X1

To study the figure, start at the bottom and move up. Note that vascular cambium (the blue cell) divides in two directions, forming secondary xylem (X) to the inside and secondary phloem (P) to the outside. These cells, which are numbered in the order in which they are produced, differentiate into the mature cell types associated with xylem and phloem. As secondary xylem accumulates, vascular cambium “moves” outward, and the woody stem increases in diameter.

P1

X1

Second division of vascular cambium forms a phloem cell. Division of vascular cambium forms two cells, one xylem cell and one vascular cambium cell. Vascular cambium cell when secondary growth begins.

Vascular cambium cell

Periderm and remnants of primary tissues

Secondary phloem

Vascular cambium Ray

Garry DeLong/Photo Researchers, Inc.

▲

FIGURE 35-5 LM of portion of a three-year-old basswood (Tilia americana) stem in cross section

Secondary xylem (3 years’ growth)

Summerwood Springwood

100 μm

Remnants of primary tissues

secondary phloem to the secondary xylem, and of waste products to the center, or heart, of the tree (discussed later).

Lenticel

Cork cambium produces periderm

James Mauseth, University of Texas

Cork cambium, which usually arises from parenchyma cells in the outer cortex, produces periderm, the functional replacement for the epidermis. Cork cambium is either a continuous cylinder of dividing cells (similar to vascular cambium) or a series of overlapping arcs of meristematic cells that form from parenchyma cells in succesCork cells sively deeper layers of the cortex and, eventually, secondary phloem. Variation in cork cambia and their rates of division explain Cork cambium and cork parenchyma why the outer bark of some tree species is fissured (as in bur oak), rough and shaggy 200 μm (shagbark hickory), scaly (Norway pine), or smooth and peeling (paper birch). As is true of vascular cambium, cork FIGURE 35-6 LM of stem periderm, showing a lenticel The epidermis has ruptured because of the proliferation of loosely arranged cork cambium divides to form new tissues in two cells in the lenticel. From the bark of a calico flower (Aristolochia elegans) stem. directions—to its inside and its outside. Cork cells, formed to the outside of cork cambium, are dead at maturity and have walls that contain layers of suberin and waxes, making them waterproof. These cork as shown in FIGURE 35-7. Bud scales cover the terminal bud and cells protect the woody stem against mechanical injury, mild fires, protect its delicate apical meristem during dormancy. When the attacks by insects and fungi, temperature extremes, and water loss. bud resumes growth, the bud scales covering the terminal bud fall To its inside, cork cambium sometimes forms cork parenchyma cells off, leaving bud scale scars on the stem where they were attached. that store water and starch granules. Cork parenchyma is only one to Because temperate-zone woody plants form terminal buds at the several cells thick, much thinner than the cork cell layer. end of each year’s growing season, the number of sets of bud scale Cork cells are impermeable to water and gases, yet the living scars on a twig indicates its age. A leaf scar shows where each leaf internal cells of the woody stem require oxygen and must be able was attached on the stem; the pattern of leaf scars can be used to to exchange gases with the surrounding atmosphere. As a stem determine leaf arrangement on a stem—alternate, opposite, or thickens from secondary growth, the epidermis, including stowhorled (see Fig. 34-2b). The vascular (conducting) tissue that mata that exchange gases for the herbaceous stem, dies. Stomata extends from the stem out into the leaf forms bundle scars within are replaced by lenticels, which permit gas exchange through the a leaf scar. Axillary buds may be found above the leaf scars. Also, periderm (FIG. 35-6). the bark of a woody twig has lenticels, which look like tiny specks on the bark of a twig. Woody twigs of deciduous trees bear characteristic features in winter Common terms associated with wood are based on plant structure A woody twig is that part of a branch produced during the current growing season. Twigs have buds, which are embryonic shoots. A If you have ever examined different types of lumber, you may terminal bud is the embryonic shoot located at the tip of a stem. have noticed that some trees have wood with two different colThe dormant (not actively growing) apical meristem of a termiors (FIG. 35-8). The functional secondary xylem—the part that nal bud is covered and protected by an outer layer of bud scales, conducts water and dissolved minerals—is the sapwood, a thin which are modified leaves (see Fig. 34-14c). Axillary buds, also layer of younger, lighter-colored wood that is closest to the bark. called lateral buds, are located in the axils of a plant’s leaves (see Heartwood, the older wood in the center of the tree, is typically a Fig. 34-1). An axil is the upper angle between a leaf and the stem brownish red. A microscopic examination of heartwood reveals to which it is attached. When terminal and axillary buds grow, they that its vessels and tracheids are plugged with pigments, tannins, form branches that bear leaves and/or flowers. The area on a stem gums, resins, and other materials. Therefore, heartwood no lonwhere each leaf is attached is called a node, and the region beger functions in conduction but instead functions as a storage site tween two successive nodes is an internode. for waste products. Because it is denser than sapwood, heartwood To demonstrate certain structural features of the stem, we can provides structural support for trees. Some evidence suggests that use a woody branch of a deciduous tree that has shed its leaves, heartwood is also more resistant to decay.

Bud scale

Terminal bud Heartwood

Sapwood

One year's growth

Axillary bud Leaf scar

Node

Carlyn Iverson

Terminal bud scale scars

Internode Node

FIGURE 35-8 Heartwood and sapwood The wood of older trees consists of a dense central heartwood and an outer layer of sapwood. The sapwood is the functioning xylem that conducts water and dissolved minerals. The annual rings in the heartwood are very conspicuous.

Lenticels Terminal bud scale scars

Bundle scars

FIGURE 35-7 Animated External structure of a woody branch in its winter condition The age of a woody branch can be determined by the number of sets of bud scale scars (do not count side branches). How old is this branch? Can you point out any twigs?

Almost everyone has heard of hardwood and softwood. Botanically speaking, hardwood is the wood of flowering plants and softwood is the wood of conifers (cone-bearing gymnosperms). The wood of pine and other conifers typically lacks fibers (with their thick secondary cell walls) and vessel elements; the conducting cells in gymnosperms are tracheids. These cell differences generally make conifer wood softer than the wood of flowering plants, although there is a substantial variation from one species to another. The balsa tree, for example, is a flowering plant whose extremely light, soft wood is used to fashion airplane models. Woody plants that grow in temperate climates where there is a growing period (during spring and summer) and a dormant period (during winter) exhibit annual rings, concentric circles found in cross sections of wood. To determine the age of a woody stem in the temperate zone, simply count the annual rings. In the tropics, environmental conditions, particularly seasonal or year-round precipitation patterns, determine the presence or absence of rings, so rings are not a reliable method of determining the ages of most tropical trees.

Examination of annual rings with a magnifying lens reveals no actual “ring,” or line, separating one year’s growth from the next. The appearance of a ring in cross section is due to differences in cell size and cell wall thickness between secondary xylem formed at the end of the preceding year’s growth and that formed at the beginning of the following year’s growth. In the spring, when water is plentiful, wood formed by vascular cambium has large-diameter conducting cells (tracheids and vessel elements) and few fibers and is appropriately called springwood or early wood (see Fig. 35-5). As summer progresses and water becomes less plentiful, the wood formed, known as summerwood or late wood, has narrower conducting cells and many fibers. It is this difference in cell size between the summerwood of one year and the springwood of the following year that gives the appearance of rings. Tree-ring analysis provides useful scientific information In temperate climates, the number of annual rings indicates the age of a tree. The size of each ring varies depending on local weather conditions. Sometimes the variation can be attributed to a single environmental factor, and similar patterns appear in the rings of different species over a large geographic area. For example, years with adequate precipitation produce wider growth rings, and years of drought produce narrower rings. It is possible to study ring sequences that are several thousand years old by constructing a master chronology, a complete sample of rings dating back as far as possible (FIG. 35-9). Dendrochronology, the study of both visible and microscopic details of tree rings, has been used extensively in several fields. Tree-ring analysis has helped date Native American prehistoric sites in the Southwest. For example, using tree-ring analysis, scientists determined that the Cliff Palace in Mesa Verde National Park dates back to the year 1073. Tree-ring analysis is also useful in ecology (to study changes in a forest community over time),

RESEARCH METHOD Why Is It Used?

By matching the rings of a wood sample of unknown age to the master chronology, investigators can accurately determine the age of the sample. Tree-ring analysis provides useful information in a variety of fields, such as to date prehistoric sites, study past climate patterns, and date major prehistoric earthquakes.

Annual rings Bark

Tilia (basswood) Long, slender core of wood extracted by a boring tool Outer bark

Vascular cambium

Annual rings

Pith

1 Sample from a living tree

1950 Outermost ring is the year when the tree was cut.

1940

1932 2 Sample from a dead tree

in the same forest

1940

1932 1931

1926

Matching and overlapping older and older wood sections extends dates back in time.

3 Sample from an old building

in the same area as the forest

1931 How Is It Done?

1926

1920

1918

A master chronology is developed using progressively older pieces of wood from the same geographic area. A small core of wood is bored out of the trunk of a living tree. The oldest rings (toward the center of the tree) are matched with the youngest rings (toward the outside) of an older tree, perhaps a dead one in the forest. The master chronology is extended back in time by overlapping matching ring sequences of successively older and older sections of wood, even those found in prehistoric dwellings. Currently, the longest master chronology—of bristlecone pines in the western United States—goes back almost 9000 years. Ring matching, once a tedious, painstaking job, is performed by computers today.

FIGURE 35-9 How tree rings are analyzed

environmental science (to study the effects of air pollution on tree growth), and geology (to date earthquakes and volcanic eruptions). Climatologists are increasingly using tree-ring data to study past climate patterns. Annual rings of certain tree species that grow at high elevations are sensitive to yearly temperature variations; the rings of these trees are wider in warm years and narrower in

cool years. Studying tree rings over long periods helps researchers determine the natural pattern of global temperature fluctuations. This information is particularly important because of concerns about the human influence on global climate. Although scientists generally agree that Earth has warmed in recent decades, they are not sure how much of the recent warming

is the result of human influence rather than natural climate variability. To help answer this vital question, European scientists are currently developing a 10,000-year master chronology that will help reconstruct annual temperatures across northern Europe and Asia since the end of the last Ice Age.

Most water that a plant absorbs is transpired through stomata into the atmosphere.

Review ■

■

■

■ ■

How does the arrangement of vascular tissue differ in primary eudicot stems and monocot stems in cross section? What is the difference between vascular cambium and cork cambium? What tissues arise from vascular cambium? from cork cambium? What happens to the primary tissues of a stem when secondary growth occurs? How do growth rings form in woody stems? What is the difference between terminal and axillary buds?

Sugar molecules manufactured in leaves by photosynthesis are transported in phloem sap throughout plant, including into roots.

Once inside roots, water and minerals are transported upward in xylem to stems, leaves, flowers, fruits, and seeds.

35.2 WATER TRANSPORT

Roots obtain water and dissolved minerals from soil.

■ ■ LEARNING OBJECTIVES 4 Describe the pathway of water movement in plants. 5 Define water potential. 6 Explain the roles of tension–cohesion and root pressure as mechanisms responsible for the rise of water and dissolved minerals in xylem.

Now that we have discussed stem structure and primary and secondary growth, we examine internal transport in the vascular system of the plant (FIG. 35-10). Roots obtain water and dissolved minerals from the soil. Once inside roots, these materials are transported upward to stems, leaves, flowers, fruits, and seeds. Furthermore, sugar molecules are transported from their place of manufacture or of breakdown of starch throughout the plant body, including into the subterranean roots. Water and dissolved minerals are transported from roots to other parts of the plant in xylem, whereas dissolved sugar is translocated in phloem. Xylem transport and phloem translocation do not resemble the movement of materials in animals because in plants nothing circulates in a system of vessels. Water and minerals, transported in xylem, travel in one direction only (upward), whereas translocation of dissolved sugar may occur upward or downward in separate phloem cells. In addition, xylem transport and phloem translocation differ from internal circulation in animals because movement in both xylem and phloem is driven largely by natural physical processes rather than by a pumping organ, or heart. How do materials travel in the continuous system of the plant’s vascular tissues? We first examine water and its movement through the plant, and later we discuss the translocation of dissolved sugar.

Water and minerals are transported in xylem Water initially moves horizontally into roots from the soil, passing through several tissues until it reaches xylem. Once the water moves into the tracheids and vessel elements of root xylem, it trav-

FIGURE 35-10 Animated Overview of transport in vascular plants Xylem transport is explained on the right side of the figure (start at the bottom and work upward), whereas phloem transport is on the left side.

els upward through a continuous network of these hollow, dead cells from root to stem to leaf. Dissolved minerals are carried along passively in the water. The plant does not expend any energy of its own to transport water, which moves as a result of natural physical processes. The transport of xylem sap is the most rapid of any movement of materials in plants (TABLE 35-1).

TABLE 35-1

Xylem and Phloem Transport Rates in Selected Plants

Plant Conifer Woody eudicot Herbaceous eudicot or monocot Herbaceous vine

Maximum Rate in Xylem (cm/min)

Maximum Rate in Phloem (cm/min)

2 73 100

0.8 2 2.8–11

250

1.2

Source: Adapted from Mauseth, J. D., Botany: An Introduction to Plant Biology, 2nd ed. Saunders College Publishing, Philadelphia, 1995. Note: Xylem and phloem rates are from different plants within each general group and should be used for comparative purposes only.

How does water move to the tops of plants? It is either pushed up from the bottom of the plant or pulled up by the top of the plant. Although plants use both mechanisms, current evidence indicates that most water is transported through xylem by being pulled to the top of the plant.

is more negative than the water potential of the soil. Thus, water moves by osmosis from the soil into the root.

Water movement can be explained by a difference in water potential

In 1896, Irish botanist Henry Dixon proposed the tension– cohesion model to explain the ascent of water against the force of gravity. Although initially met with widespread skepticism in the scientific community, this model has withstood the test of time. According to the tension–cohesion model, also known as the transpiration–cohesion model, water is pulled up the plant as a result of a tension produced at the top of the plant (FIG. 35-11). This tension, which resembles that produced when drinking a liquid through a straw, is caused by the evaporative pull of transpiration. Recall from Chapter 34 that transpiration is the evaporation of water vapor from plants. Most water loss from transpiration takes place through stomata, the numerous microscopic pores present on leaf and stem surfaces. Water vapor diffuses through the stomata into a boundary layer of quiet air adjacent to the leaf surface. The boundary layer is thicker when the wind is calm than when it is gusty; a thicker boundary layer results in a slower rate of transpiration. The tension extends from leaves, where most transpiration occurs, down the stems and into the roots. It draws water up stem xylem to leaf cells that have lost water as a result of transpiration and pulls water from root xylem into stem xylem. As water is pulled upward, additional water from the soil is drawn into the roots. Thus, the pathway of water movement is as follows:

To understand how water moves, it is helpful to introduce water potential, which is defined as the free energy of water (see Chapter 7). Water potential (Cw), represented by the Greek letter psi, is important in plant physiology because it is a measure of a cell’s ability to absorb water by osmosis. (Recall from Chapter 5 and Fig. 5-12 that osmosis is a special kind of diffusion that involves the net movement of water through a selectively permeable membrane from a region of higher concentration to a region of lower concentration.) Water potential also provides a measure of water’s tendency to evaporate from cells. The water potential of pure water is conventionally set at 0 megapascals (MPa) because it cannot be measured directly. (A megapascal is a unit of pressure equal to about 10.2 kg/cm2, or 145.1 lb/in2.) However, botanists can measure differences in the free energy of water molecules in different situations. When solutes dissolve in water, the free energy of water decreases. Solutes induce hydration, in which water molecules surround ions and polar molecules, keeping them in solution by preventing them from coming together (see Fig. 2-10). The association of water molecules with hydrated molecules and ions reduces the motion of water molecules, decreasing their free energy. Thus, dissolved solutes lower the water potential to a negative number. Water moves from a region of higher (less negative) water potential to a region of lower (more negative) water potential. The water potential of the soil varies, depending on how much water it contains. When a soil is extremely dry, its water potential is very low (very negative). When a soil is moister, its water potential is higher, although it still has a negative value because dissolved minerals are present in dilute concentrations (TABLE 35-2). The water potential in root cells is also negative because of the presence of dissolved solutes. Roots contain more dissolved materials than soil water does, unless the soil is extremely dry. This means that under normal conditions the water potential of the root

TABLE 35-2

Representative Water Potential (CW) Values*

Plant Parts and the Nearby Environment Atmosphere near leaves Leaf interior Stem xylem Root interior Soil water

Water Potentials in Megapascals (MPa) −80.0 −1.5 −0.7 −0.4 −0.1

*Actual values vary widely, depending on environmental factors such as solute concentration, hydrostatic pressure, and gravity.

According to the tension–cohesion model, water is pulled up a stem

soil ¡ root tissues (epidermis, cortex, and so forth) ¡ root xylem ¡ stem xylem ¡ leaf xylem ¡ leaf mesophyll ¡ atmosphere

This upward pulling of water is possible only as long as there is an unbroken column of water in xylem throughout the plant. Water forms an unbroken column in xylem because of the cohesiveness of water molecules. Recall from Chapter 2 that water molecules are cohesive, that is, strongly attracted to one another, because of hydrogen bonding. In addition, the adhesion of water to the walls of xylem cells, also the result of hydrogen bonding, is an important factor in maintaining an unbroken column of water. Thus, the cohesive and adhesive properties of water enable it to form a continuous column that can be pulled up through the xylem. The movement of water in xylem by the tension–cohesion mechanism can be explained in terms of water potential. The atmosphere has an extremely negative water potential. For example, air with a relative humidity of 50% has a water potential of −100 MPa; even moist air at a relative humidity of 90% has a negative water potential of −13 MPa. Thus, there is a water potential gradient from the least negative (the soil) up through the plant to the most negative (the atmosphere). This gradient literally pulls the water from the soil up through the plant. Although the tension–cohesion model was first proposed toward the end of the 19th century, conclusive experimental evidence to support this mechanism was not obtained until the late

KEY POINT

Transpiration is the driving force of the tension–cohesion model.

Leaf xylem Mesophyll cells Stoma 1

Water molecules

Stem xylem

2

3

Water vapor transpires from the surfaces of leaf mesophyll cells to the drier atmosphere through stomata. This produces a tension that pulls water out of leaf xylem toward mesophyll cells.

Cohesion of water molecules, caused by hydrogen bonding, allows unbroken columns of water to be pulled up narrow vessels and tracheids of stem xylem.

This, in turn, pulls water up root xylem, forming a continuous column of water from root xylem to stem xylem to leaf xylem. Movement of water upward in root produces a pull that causes soil water to diffuse into the root.

Root xylem

Soil water Root hair

FIGURE 35-11 Animated The tension–cohesion model

1990s and early 2000s. At that time, several research groups made direct measurements of the large negative pressure that exists in xylem, indicating that the water potential gradients in root, stem, and leaf xylem are adequate to explain the observed movement of water.

Is the tension–cohesion model powerful enough to explain the rise of water in the tallest plants? Plant biologists have studied this question for years. For example, a 2004 study of five of the eight tallest trees in the world (all redwoods, or Sequoia sempervirens) concluded that the tension produced by transpiration is strong enough

to pull water upward to a maximum height of 130 m (422 ft). Because the height of the tallest known living tree, a coast redwood in California, measured 117 m (379 ft) in 2007, the tension–cohesion model easily accounts for the transport of water. Botanists generally consider the tension–cohesion model to be the dominant mechanism of xylem transport in most plants.

Root pressure pushes water from the root up a stem In the less important mechanism for water transport, known as root pressure, water that moves into roots from the soil is pushed up through xylem toward the top of the plant. Root pressure occurs because mineral ions that are actively absorbed from the soil are pumped into the xylem, decreasing its water potential. This accumulation of ions has an osmotic effect, causing water to move into xylem cells from surrounding root cells. In turn, water moves into roots by osmosis because of the difference in water potential between the soil and root cells. The accumulation of water in root tissues produces a positive pressure (as high as +0.2 MPa) that forces the water up through the root xylem into the shoot. Guttation, a phenomenon in which liquid water is forced out through special openings in the leaves (see Fig. 34-12), results from root pressure. However, root pressure is not strong enough to explain the rise of water to the tops of tall trees. Root pressure exerts an influence in much smaller plants, particularly in the spring when the soil is moist, but it clearly does not cause water to rise to the tops of the tallest plants. Furthermore, root pressure does not occur to any appreciable extent in summer (when water is often not plentiful in soil), yet water transport is greatest during hot summer days.

Review ■ ■ ■

What is water potential? How is the movement of water related to water potential? How does the tension–cohesion model explain the rise of water in the tallest trees?

35.3 TRANSLOCATION OF SUGAR IN SOLUTION ■ ■ LEARNING OBJECTIVES 7 8

Describe the pathway of sugar translocation in plants. Discuss the pressure–flow model of sugar translocation in phloem.

The sugar produced during photosynthesis is converted into sucrose (common table sugar), a disaccharide composed of one molecule of glucose and one of fructose (see Fig. 3-8b), before being loaded into phloem and translocated to the rest of the plant. Sucrose is the predominant photosynthetic product carried in phloem. Phloem sap also contains much smaller amounts of other materials, such as amino acids, organic acids, proteins, hormones,

and certain minerals (such as potassium, chloride, phosphate, and magnesium). Sometimes disease-causing plant viruses are transported throughout the plant in phloem. Translocation of phloem sap is not as rapid as xylem transport (see Table 35-1). Fluid within phloem tissue moves both upward and downward. Sucrose is translocated in individual sieve tubes from a source, an area of excess sugar supply (usually a leaf), to a sink, an area of storage (as insoluble starch) or of sugar use, such as roots, apical meristems, fruits, and seeds.

The pressure–flow model explains translocation in phloem Current experimental evidence supports the translocation of dissolved sugar in phloem by the pressure–flow model, which was first proposed in 1926 by the German scientist Ernst Münch. The pressure–flow model states that solutes (such as dissolved sugars) move in phloem by means of a pressure gradient—that is, a difference in pressure. The pressure gradient exists between the source, where the sugar is loaded into phloem, and the sink, where the sugar is removed from phloem. At the source, the dissolved sucrose is moved from a leaf ’s mesophyll cells, where it was manufactured, into the companion cells, which load sucrose into the sieve tube elements of phloem. Sucrose loading occurs by active transport, a process that requires adenosine triphosphate (ATP) (FIG. 35-12). The ATP supplies energy to pump protons (H+) out of the sieve tube elements, producing a proton gradient that drives the uptake of sugar through specific channels by the cotransport of protons back into the sieve tube elements. (Recall the discussion of a cotransport system in Chapter 5.) In the cotransport system involved in phloem loading, sugar is moved from a region of low concentration to a region of high concentration by coupling its transport to the transport of protons down their concentration gradient. The sugar therefore accumulates in the sieve tube element. The increase in dissolved sugars in the sieve tube element at the source—a concentration that is two to three times as great as in surrounding cells—decreases (makes more negative) the water potential of that cell. As a result, water moves by osmosis from nearby xylem cells into the sieve tubes, increasing the turgor pressure (hydrostatic pressure) inside them. Thus, phloem loading at the source occurs as follows: proton pump moves H+ out of sieve tube element ¡ sugar is actively transported into sieve tube element ¡ water diffuses from xylem into sieve tube element ¡ turgor pressure increases within sieve tube

At its destination (the sink), sugar is unloaded by various mechanisms, both active and passive, from the sieve tube elements. With the loss of sugar, the water potential in the sieve tube elements at the sink increases (becomes less negative). Therefore, water moves out of the sieve tubes by osmosis and into surrounding cells where the water potential is more negative. Most of this water diffuses back to the xylem to be transported upward. This water movement de-

The pressure–flow model explains the movement of dissolved sugar in phloem by means of a pressure gradient. The difference in sugar concentrations between the source and the sink causes translocation in phloem XYLEM PHLOEM as water and dissolved sugar flow along the 1 Sucrose is actively pressure gradient. This pressure gradient loaded into sieve tube pushes the sugar solution through phloem elements (requires much as water is forced through a hose. ATP). SOURCE The actual translocation of dissolved Leaf cell sugar in phloem does not require metabolic 2 Water diffuses energy. However, the loading of sugar at the 2 from xylem as a result source and the active unloading of sugar at 1 of decreased (more the sink require energy derived from ATP to negative) water move the sugar across cell membranes by acpotential in sieve tube. tive transport. Although the pressure–flow model adequately explains current data on phloem 3 The increase in pressure forces the translocation, much remains to be learned 3 fluid through the sieve Companion cells about this complex process. Phloem translotube to the root. cation is difficult to study in plants. Because Sieve tube element phloem cells are under pressure, cutting into phloem to observe it releases the pressure 4 Sucrose is actively Direction of and causes the contents of the sieve tube and passively unloaded water movement elements (the phloem sap) to exude and into sink cell, such as Direction of parenchyma cell in the mix with the contents of other severed cells sucrose movement root cortex. (Active that are also unavoidably cut. In the 1950s, unloading requires SINK 6 scientists developed a unique research tool Root cell ATP.) to avoid contaminating the phloem sap: aphids, which are small insects that insert their mouthparts into phloem sieve tubes 5 Water diffuses for feeding (FIG. 35-13). The pressure in the from phloem to xylem 4 as a result of increased punctured phloem drives the sugar solution 5 (less negative) water through the aphid’s mouthpart into its dipotential in sieve tube. gestive system. When the aphid’s mouthpart is severed from its body by a laser beam, the sugar solution continues to flow through the 6 Once water is in the mouthpart at a rate proportional to the presxylem, transpiration sure in phloem. This rate can be measured, pulls water to leaves and the effects on phloem transport of difagain. ferent environmental conditions—varying Vessel Sieve tube light intensities, darkness, and mineral defirunning through running through ciencies, for example—can be ascertained. length of plant length of plant The identity and proportions of translocated substances can also be determined FIGURE 35-12 Animated The pressure–flow model using severed aphid mouthparts. This technique has verified that in most plant species the sugar sucrose is the primary carbohydrate transported in phloem; however, some species transport creases the turgor pressure inside the sieve tubes at the sink. Thus, other sugars, such as raffinose, or sugar alcohols, such as sorbitol. phloem unloading at the sink proceeds as follows:

KEY POINT

In phloem, solutes move from sources to sinks. The pressure gradient within the sieve tube causes translocation from the area of higher turgor pressure (the source) to the area of lower turgor pressure (the sink).

Review sugar is transported out of sieve tube element ¡ water diffuses out of sieve tube element and into xylem ¡ turgor pressure decreases within sieve tube

■ ■

How does the direction of transport differ in xylem and phloem? How does the pressure–flow model explain sugar movement in phloem? Include in your answer the activities at source and sink.

KEY EXPERIMENT QUESTION: How can phloem sap be studied without cutting nonphloem cells that would contaminate the sap? HYPOTHESIS: An aphid mouthpart can be used to penetrate a single sieve tube.

Mouthpart Sieve tube element

Dwight Kuhn

Mouthpart

M. H. Zimmermann, SCIENCE, Vol. 133, pp. 73–79 (Fig. 4), 13 Jan. 1961. © 1961 by the American Association for the Advancement of Science

EXPERIMENT: After allowing aphids to insert their mouthparts into phloem of a stem, researchers anesthetized the feeding aphids with CO2 and used a laser to cut their bodies away from their mouthparts. The mouthparts remained in the phloem and functioned like miniature pipes.

25 μm

(a) Aphid feeding on a stem.

(b) LM of phloem cells, showing an aphid mouthpart penetrating a sieve tube element.

RESULTS AND CONCLUSION: About 1 mm3 of phloem sap exuded from each severed mouthpart per hour for several days, so researchers were able to collect and analyze the composition of the sap.

FIGURE 35-13 Collecting and analyzing phloem sap This method, first used in the 1950s by insect physiologists studying aphids, was quickly adopted by plant physiologists studying phloem translocation.

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

35.1 (page 745) 1 Label cross sections of herbaceous eudicot and monocot stems, and describe the functions of each tissue. Herbaceous stems possess an epidermis, vascular tissue, and either ground tissue or cortex and pith. The epidermis is a protective layer covered by a water-conserving cuticle. Stomata permit gas exchange. Xylem conducts water and dissolved minerals, and phloem conducts dissolved sugar. The cortex, pith, and ground tissue function primarily for storage. ■ All herbaceous stems have the same basic tissues, but their arrangement varies. Herbaceous eudicot stems have the vascular bundles arranged in a circle (in cross section) and have a distinct cortex and pith. Monocot stems have vascular bundles scattered in ground tissue. ■

Learn more about eudicot and monocot stems by clicking on the figures in CengageNOW.

2 Name the two lateral meristems and describe the tissues that arise from each. ■ Vascular cambium is the lateral meristem that produces secondary xylem (wood) and secondary phloem (inner bark). ■ Cork cambium produces periderm, which consists of cork parenchyma and cork cells. Cork cells are the functional replacement for epidermis in a woody stem. Cork parenchyma functions primarily for storage in a woody stem. 3 Outline the transition from primary growth to secondary growth in a woody stem. Secondary growth (the production of the secondary tissues, wood ■ and bark) occurs in some flowering plants (woody eudicots) and in all cone-bearing gymnosperms. During secondary growth, the vascular cambium divides in two directions to form secondary xylem (to the inside) and secondary phloem (to the outside). The primary xylem and primary phloem in the original vascular bundles become separated as secondary growth proceeds.

Learn more about the tension–cohesion model by clicking on the figure in CengageNOW. ■

Root pressure, caused by the movement of water into roots from the soil as a result of the active absorption of mineral ions from the soil, helps explain the rise of water in smaller plants, particularly when the soil is wet. Root pressure pushes water up through xylem.

35.3 (page 756) 7 Describe the pathway of sugar translocation in plants. Dissolved sugar is translocated upward or downward in phloem, from a source (an area of excess sugar, usually a leaf) to a sink (an area of storage or of sugar use, such as roots, apical meristems, fruits, and seeds). Sucrose is the predominant sugar translocated in phloem. 8 Discuss the pressure–flow model of sugar translocation in phloem. ■ Movement of materials in phloem is explained by the pressure– flow model. Companion cells actively load sugar into the sieve tubes at the source; ATP is required for this process. The ATP supplies energy to pump protons out of the sieve tube elements. The proton gradient drives the uptake of sugar by the cotransport of protons back into the sieve tube elements. Sugar therefore accumulates in the sieve tube element, causing the movement of water into the sieve tubes by osmosis. ■ Companion cells actively (requiring ATP) and passively (not requiring ATP) unload sugar from the sieve tubes at the sink. As a result, water leaves the sieve tubes by osmosis, decreasing the turgor pressure (hydrostatic pressure) inside the sieve tubes. ■ The flow of materials between source and sink is driven by the turgor pressure gradient produced by water entering phloem at the source and water leaving phloem at the sink. ■

Learn more about secondary growth by clicking on the figures in CengageNOW.

35.2 (page 753) 4 Describe the pathway of water movement in plants. Water and dissolved minerals move from the soil into root tissues (epidermis, cortex, and so forth). Once in root xylem, water and minerals move upward, from root xylem to stem xylem to leaf xylem. Much of the water entering the leaf exits leaf veins and passes into the atmosphere. 5 Define water potential. ■ Water potential (Cw) is a measure of the free energy of water. Pure water has a water potential of 0 megapascals (MPa), whereas water with dissolved solutes has a negative water potential. Water moves from an area of higher (less negative) water potential to an area of lower (more negative) water potential. ■

6 Explain the roles of tension–cohesion and root pressure as mechanisms responsible for the rise of water and dissolved minerals in xylem. ■ The tension–cohesion model explains the rise of water in even the tallest plants. The evaporative pull of transpiration causes tension at the top of the plant. This tension is the result of a water potential gradient that ranges from the slightly negative water potentials in the soil and roots to the very negative water potentials in the atmosphere. As a result of the cohesive and adhesive properties of water, the column of water pulled up through the plant remains unbroken.

Learn more about how sugar travels by clicking on the figure in CengageNOW.

T E S T YO U R U N D E R S TA N D I N G 1. Axillary buds are located (a) at the tips of stems (b) in unusual places, such as on roots (c) in the region between two successive nodes (d) in the upper angle between a leaf and the stem to which it is attached (e) within the loosely arranged cells of lenticels 2. Ground tissue in monocot stems performs the same functions as _____ and _____ in herbaceous eudicot stems. (a) phloem; xylem (b) cork cambium; vascular cambium (c) epidermis; periderm (d) primary xylem; secondary xylem (e) cortex; pith 3. Which of the following statements is false? (a) primary growth is an increase in the length of a plant (b) primary growth occurs at both apical and lateral meristems (c) all plants have primary growth (d) herbaceous stems have primary growth, whereas woody stems have both primary and secondary growth (e) buds are embryonic shoots that contain apical meristems 4. The two lateral meristems responsible for secondary growth are (a) phloem and xylem (b) cork cambium and vascular cambium (c) epidermis and periderm (d) primary xylem and secondary xylem (e) cortex and pith

5. Cork cambium and the tissues it produces are collectively called (a) periderm (b) lenticels (c) cortex (d) epidermis (e) wood 6. Each annual ring in a section of wood represents one year’s growth of (a) primary xylem (b) secondary xylem (c) primary xylem or secondary xylem in alternate years (d) primary phloem (e) secondary phloem 7. Water potential is (a) the formation of a proton gradient across a cell membrane (b) the transport of a watery solution of sugar in phloem (c) the transport of water in both xylem and phloem (d) the removal of sucrose at the sink, causing water to move out of the sieve tubes (e) the free energy of water in a particular situation 8. Which of the following is a mechanism of water movement in xylem that is responsible for guttation? (a) pressure–flow (b) tension–cohesion (c) root pressure (d) active transport of potassium ions into guard cells (e) transpiration 9. Which of the following is a mechanism of water movement in xylem that combines the evaporative pull of transpiration with

the cohesive and adhesive properties of water? (a) pressure–flow (b) tension–cohesion (c) root pressure (d) active transport of potassium ions into guard cells (e) guttation 10. Which of the following is a mechanism of phloem transport in which dissolved sugar is moved by means of a pressure gradient that exists between the source and the sink? (a) pressure–flow (b) tension–cohesion (c) root pressure (d) active transport of potassium ions into guard cells (e) guttation

becomes more negative (c) water potential becomes more positive under certain conditions and more negative under other conditions (d) water potential is not affected by solute concentration (e) water potential is always zero when solutes are dissolved in water 12. Label the various tissues, give at least one function for each tissue, and identify the stem as a herbaceous eudicot, monocot, or woody plant. Use Figure 35-1 to check your answers.

Ed Reschke/Peter Arnold, Inc.

Ed Reschke/Peter Arnold, Inc.

11. How does increasing solute concentration affect water potential? (a) water potential becomes more positive (b) water potential

(a)

(b)

CRITICAL THINKING 1. When secondary growth is initiated, certain cells become meristematic and begin to divide. Could a mature tracheid ever do this? a sieve tube element? Why or why not? 2. Why does the wood of many tropical trees lack annual rings? Why does the wood of other tropical trees possess annual rings? 3. Why is hardwood more desirable than softwood for making furniture? Explain your answer, based on the structural differences between hardwood and softwood. 4. Why should you cut off a few inches from the stem ends of cut flowers before placing them in water? Base your answer on what you have learned about the tension–cohesion model. 5. When a strip of bark is peeled off a tree branch, what tissues are usually removed? 6. Could a tree grow to a height of 150 m (488 ft)? Why or why not?

7. EVOLUTION LINK. Like stems in general, some vines are herbaceous and others are woody. Tropical rain forests have a greater diversity of vines than in any other environment on Earth, and most of these vines are woody. Develop a hypothesis to explain why natural selection has favored the evolution of more species of woody vines (as opposed to herbaceous vines) in tropical rain forests. 8. ANALYZING DATA. A plant cell with a water potential Cw = −1.5 MPa is adjacent to a second cell with Cw = −1.8 MPa. Will water flow by osmosis from one cell to the other? If so, in which direction? Additional questions are available in CengageNOW at www.cengage.com/ login.

36

Roots and Mineral Nutrition

© Inga Spence/Alamy

Storage roots. Cassava (Manihot esculenta) plants have been dug up to show the growth habit of the storage roots. Cassava roots are an important staple crop in the tropics. Some cassava roots contain a poison (prussic acid), which is destroyed during cooking.

KEY CONCEPTS 36.1 Root structure is related to function. Primary tissues (epidermis, cortex, endodermis, pericycle, xylem, and phloem) of roots develop from root apical meristems. Secondary tissues (wood and bark) of woody roots develop from lateral meristems. 36.2 Roots associate and interact with many organisms, including soil bacteria and fungi.

36.3 Soil, the layer of Earth’s crust that has been modified by contact with weather, wind, water, and organisms, contains most of the essential elements required by plants.

B

ranching underground root systems are often more extensive than a plant’s aerial parts. The roots of a corn plant, for example, may grow

to a depth of 2.5 m (about 8 ft) and spread outward 1.2 m (4 ft) from the stem. Desert-dwelling tamarisk (Tamarix) trees reportedly have roots that grow to a depth of 50 m (163 ft) to tap underground water. The extent of a plant’s root depth and spread varies considerably among different species and even among different individuals in the same species. Because roots are usually underground and out of sight, people do not always appreciate the important functions they perform. First, as anyone who has ever pulled weeds can attest, roots anchor a plant securely in the soil. A plant needs a solid foundation from which to grow. Firm anchorage is essential to a plant’s survival so that the stem remains upright, enabling leaves to absorb sunlight effectively. Second, roots absorb water and dissolved minerals (inorganic nutrients) such as nitrates, phosphates, and sulfates, which are necessary for synthesizing important organic molecules. These dissolved minerals are then transported throughout the plant in the xylem. Many roots perform the function of storage. Carrots, sweet potatoes, cassava, and other root crops are important sources of human food (see

photograph). Surplus sugars produced in the leaves by photosynthesis are transported in the phloem to the roots for food storage (usually as starch or sucrose) until needed. Other plants, particularly those living in arid regions, possess storage roots adapted to store water. In certain species, roots are modified for functions other than anchorage, absorption, conduction, and storage. Roots specialized to perform uncommon functions are discussed later in the chapter.

36.1 ROOT STRUCTURE AND FUNCTION ■ ■ LEARNING OBJECTIVES 1 2 3 4 5

Distinguish between taproot and fibrous root systems. Compare cross sections of a primary eudicot root and a monocot root, and describe the functions of each tissue. Trace the pathway of water and mineral ions from the soil through the various root tissues, and distinguish between the symplast and apoplast. Discuss the structure of roots with secondary growth. Describe at least three roots that are modified to perform uncommon functions.

Two types of root systems, a taproot system and a fibrous root system, occur in plants (FIG. 36-1). A taproot system consists of one main root that formed from the seedling’s enlarging radicle, or

(b)

(a)

FIGURE 36-1 Animated Root systems (a) A taproot system develops from the embryonic root in the seed. (b) The roots of a fibrous root system are adventitious and develop from stem tissue.

embryonic root. Many lateral roots of various sizes branch out of a taproot. Taproots are characteristic of many eudicots and gymnosperms. A dandelion is a good example of a common herbaceous plant with a taproot system. A few trees, such as hickory, retain their taproots, which become quite massive as the plants age. Most trees, however, have taproots when young and later develop large, shallow, lateral roots from which other roots branch off and grow downward. A fibrous root system has several to many roots of similar size developing from the end of the stem, with lateral roots branching off these roots. Fibrous root systems form in plants that have a short-lived embryonic root. The roots originate first from the base of the embryonic root and later from stem tissue. The main roots of a fibrous root system do not arise from pre-existing roots but from the stem; such roots are called adventitious roots. Adventitious organs occur in an unusual location, such as roots that develop on a stem, or buds that develop on roots. Onions, crabgrass, and other monocots have fibrous root systems. Taproot and fibrous root systems are adapted to obtain water in different sections of the soil. Taproot systems often extend down into the soil to obtain water located deep underground, whereas fibrous root systems, which are located relatively close to the soil surface, are adapted to obtain rainwater from a larger area as it drains into the soil.

Roots have root caps and root hairs Because of the need to adapt to the soil environment instead of the atmospheric environment, roots have several structures, such as root caps and root hairs, that shoots lack. Although stems and leaves have various types of hairs, they are distinct from root hairs in structure and function. Each root tip is covered by a root cap, a protective, thimblelike layer many cells thick that covers the delicate root apical meristem (FIG. 36-2a; also see Fig. 33-6). As the root grows, pushing its way through the soil, cells of the root cap are sloughed off by the frictional resistance of the soil particles and replaced by new cells formed by the root apical meristem. The root cap cells secrete lubricating polysaccharides that reduce friction as the root passes through the soil. The root cap also appears to be involved in orienting the root so that it grows downward (see discussion of gravitropism in Chapter 38). When a root cap is removed, the root apical meristem grows a new cap. However, until the root cap has regenerated, the root grows randomly rather than in the direction of gravity. Root hairs are short-lived tubular extensions of epidermal cells located just behind the growing root tip. Root hairs continually form in the area of cell maturation closest to the root tip to replace those that are dying off at the more mature end of the roothair zone (FIG. 36-2b; also see Fig. 33-6). Each root hair is short (typically less than 1 cm, or 0.4 in, in length), but they are quite numerous. Root hairs greatly increase the absorptive capacity of roots by increasing their surface area in contact with moist soil. Soil particles are coated with a microscopically thin layer of water in which minerals are dissolved. The root hairs establish an intimate contact with soil particles, which allows efficient absorption of water and minerals.

Runk/Rannels/Grant Heilman Photography

Root apical meristem (area of cell division) Root cap Root hairs Soil air

Cengage

Soil water Soil particles

250 μm

(a) Root cap. LM of an oak (Quercus sp.) root tip showing its root cap. The root apical meristem is protected by the root cap.

Epidermis

(b) Root hairs. Each delicate hair is an extension of a single cell of the root epidermis. Root hairs increase the surface area of the root in contact with the soil. This radish (Raphanus sativus ) seedling is approximately 5 cm (2 in) long.

FIGURE 36-2 Animated Structures unique to roots

The development of root hairs is under the control of two regulatory genes that code for transcription factors. (You may recall from Chapter 14 that transcription factors are DNA-binding proteins that regulate the synthesis of RNA from a DNA template.) In 2007, researchers from the United Kingdom and France reported in the journal Science that these two genes have an ancient origin with evolutionary significance. The biologists compared two genes that control rhizoid formation in the gametophyte stage of the moss Physcomitrella patens to those that control root hair formation in the sporophyte stage of the angiosperm Arabidopsis thaliana. They found that the genes code for transcription factors that are essentially the same in both plants. Mosses such as Physcomitrella represent an ancient lineage of land plants (see Chapter 27). It therefore appears that these genes were conserved during the evolution from a dominant gametophyte in mosses to a dominant sporophyte in flowering plants.

The arrangement of vascular tissues distinguishes the roots of herbaceous eudicots and monocots Although considerable variation exists in herbaceous eudicot and monocot roots, they all have an outer protective covering (epidermis), a cortex for storage of starch and other organic molecules, and vascular tissues for conduction. Let us first consider the structure of herbaceous eudicot roots. In most herbaceous eudicot roots, the central core of vascular tissue lacks pith The buttercup root is a representative eudicot root with primary growth (FIG. 36-3). Like other parts of this herbaceous eudicot, a single layer of protective tissue, the epidermis, covers its roots.

The root hairs are a modification of the root epidermis that enables it to absorb more water from the soil. The root epidermis does not secrete a thick, waxy cuticle in the region of root hairs because this layer would impede the absorption of water from the soil. Both the lack of a cuticle and the presence of root hairs increase absorption. Beginning with the root hairs, most of the water that enters the root moves along the cell walls rather than enters the cells. One of the major components of cell walls is cellulose, which absorbs water as a sponge does. An example of the absorptive properties of cellulose is found in cotton balls, which are almost pure cellulose. The cortex, which is composed primarily of loosely packed parenchyma cells, composes the bulk of a herbaceous eudicot root. Roots usually lack supporting collenchyma cells, probably because the soil supports the root, although roots as they age may develop some sclerenchyma (another supporting tissue; see Chapter 33). The primary function of the root cortex is storage. A microscopic examination of the parenchyma cells that form the cortex often reveals numerous amyloplasts (see Figs. 36-3b and 3-9a), which store starch. Starch, an insoluble carbohydrate composed of glucose subunits, is the most common form of stored energy in plants. When used at a later time, these reserves provide energy for such activities as growth and cell replacement following an injury. The large intercellular (between-cell) spaces, a common feature of the root cortex, provide a pathway for water uptake and allow for aeration of the root. The oxygen that root cells need for aerobic respiration diffuses from air spaces in the soil into the intercellular spaces of the cortex and from there into the cells of the root. The water and dissolved minerals that enter the root cortex from the epidermis move in solution along two pathways: the symplast and apoplast (FIG. 36-4). The symplast is the continuum of

Epidermis

Cortex cells filled with amyloplasts

Cortex

Endodermis cell Pericycle cell

Stele

Ed Reschke/Peter Arnold, Inc.

Ed Reschke/Peter Arnold, Inc.

Phloem cell Xylem vessel elements Intercellular space in cortex

250 μm

25 μm

(a) Eudicot root. Cortex composes the bulk of herbaceous eudicot roots. Note the X-shaped xylem in the center of the root.

(b) Stele of eudicot root. Surrounding the solid core of vascular tissues is a single layer of pericycle, which is meristematic in growing roots.

FIGURE 36-3 LMs of cross sections of a herbaceous eudicot root Shown is a buttercup (Ranunculus) root.

KEY POINT

The symplast pathway (blue) consists of living cells interconnected by plasmodesmata. The apoplast pathway (red) uses cell walls and intercellular spaces. Movement upward by tension–cohesion

Xylem vessels

Endodermis Casparian strip Cortex

Epidermis

Symplast: interconnected cytoplasm of living cells Phloem cells Pericycle

Water and dissolved nutrient minerals

Plasma membrane Plasmodesma Cell wall Apoplast: interconnected cell wall spaces

Root hair

FIGURE 36-4 Pathways of water and dissolved minerals in the root Water and dissolved minerals travel from cell to cell along the interconnected porous cell walls (the apoplast) or from one cell’s cytoplasm to another through plasmodesmata (the symplast). On reaching the

endodermis, water and minerals can move into the root’s center only if they pass through a plasma membrane and enter the cytoplasm of an endodermal cell.

living cytoplasm, which is connected from one cell to the next by cytoplasmic bridges called plasmodesmata (see Fig. 5-27). Some dissolved mineral ions move from the epidermis through the cortex via the symplast. The apoplast consists of the interconnected porous cell walls of a plant, along which water and mineral ions move freely. The water and mineral ions can diffuse across the cortex without ever crossing a plasma membrane to enter a living cell. Together the symplast and apoplast make up the entire plant body. The inner layer of the cortex, the endodermis, regulates the movement of water and minerals that enter the xylem in the root’s center. Structurally, the endodermis differs from the rest of the cortex. Endodermal cells fit snugly against one another, and each has a special bandlike region, called a Casparian strip (FIG. 36-5), on its radial (side) and transverse (upper and lower) walls. Casparian strips contain suberin, a fatty material that is waterproof. (Recall from Chapter 33 that suberin is also the waterproof material in cork cell walls.) Until the endodermis is reached, most of the water and dissolved minerals have traveled along the apoplast and therefore have not passed through a plasma membrane or entered the cytoplasm of a root cell. However, the waterproof Casparian strip on the radial and transverse walls of the endodermal cells prevents water and minerals from continuing to move passively along the cell walls. For substances to pass farther into the root interior, they must move from the cell walls into the cytoplasm of the endodermal cells. Water enters by osmosis, both directly across the lipid bilayer of the endodermal plasma membrane and through

KEY POINT

aquaporins, integral membrane proteins that facilitate the rapid transport of water across the membrane (FIG. 36-6a). As in water movement across the lipid bilayer, water moves through aquaporins only along the osmotic gradient, from a region of higher water concentration to a region of lower water concentration. The key role of aquaporins in the uptake of water by root cells is currently under active study. Minerals enter endodermal cells by passing through carrier proteins in their plasma membranes. In carrier-mediated active transport, the mineral ions are pumped against their concentration gradient—that is, from an area of low concentration of that mineral in the soil solution to an area of high concentration in the plant’s cells (FIG. 36-6b). One of many reasons why root cells require sugar and oxygen for aerobic respiration is that this active transport requires the expenditure of energy, usually in the form of ATP. From the endodermis, water and mineral ions enter the root xylem (botanists do not yet know precisely how this is done) and are conducted to the rest of the plant. At the center of a eudicot primary root is the stele, or vascular cylinder, a central cylinder of vascular tissues. The outermost layer of the stele is the pericycle, which is just inside the endodermis. The pericycle consists of a single layer of parenchyma cells that give rise to multicellular lateral roots, also called branch roots (FIG. 36-7). Lateral roots originate when cells in a portion of the pericycle start dividing. As it grows, the lateral root pushes through several layers of root tissue (endodermis, cortex, and epidermis) before entering the soil. Each lateral root has all the structures

The endodermis regulates the kinds of minerals that are absorbed from the soil and conducted to the rest of the plant body. Entry via apoplast Entry via symplast Endodermis

Cortex

Epidermis

Cortex

Casparian strip

Endodermis

Root cross section

Casparian strip

Movement of water through the endodermis to the center of the root

FIGURE 36-5 Animated Endodermis and mineral uptake Note the Casparian strip around the radial and transverse walls that prevents water and dissolved minerals from passing into the stele along endodermal cell walls. To reach the vascular tissues, water and dissolved minerals must pass through the plasma membranes of endodermal cells.

Water molecules

Outside cell

Ruptured epidermis and cortex Lateral root

Cortex

Cell wall

Lipid bilayer of plasma membrane

Aquaporin

James Mauseth, University of Texas

Cytosol

(a) Movement of water. Water crosses membranes by diffusion through the lipid layer (left) and, more rapidlly, through aquaporins (right). Outside cell

Ion transport channel

Protons 100 μm

Proton pump

FIGURE 36-7 LM of a lateral root Lateral roots originate at the pericycle.

ATP ADP

Ions

Cytosol

(b) Movement of mineral ions by carrier-mediated active transport. The lipid bilayer of plant membranes is impermeable to mineral ions, which can only pass through ion transport channels. The hydrolysis of ATP pumps protons outside the cell, forming an energy-charged proton gradient. Ion transport channels allow mineral ions to enter the cell with protons.

FIGURE 36-6 Transport of water and minerals across plant membranes

and features—root cap, root hairs, epidermis, cortex, endodermis, pericycle, xylem, and phloem—of the larger root from which it emerges. In addition to producing lateral roots, the pericycle is involved in forming the lateral meristems that produce secondary growth in woody roots (discussed later in the chapter). Xylem, the centermost tissue of the stele, often has two, three, four, or more extensions, or “xylem arms” (see Fig. 36-3b). Phloem is located in patches between the xylem arms. The xylem and phloem of the root have the same functions and kinds of cells as in the rest of the plant: water and dissolved minerals are conducted in tracheids and vessel elements of xylem, and dissolved sugar (sucrose) is conducted in sieve tube elements of phloem.

After passing through the endodermal cells, water enters the root xylem, often at one of the xylem arms. Up to this point, the pathway of water has been horizontal from the soil into the center of the root: root hair/epidermis ¡ cortex ¡ endodermis ¡ pericycle ¡ root xylem

Once water enters the xylem, it is transported upward through root xylem into stem xylem and from there to the rest of the plant. One direction of phloem conduction is from the leaves, where sugar is made by photosynthesis, to the root, where sugar is used for the growth and maintenance of root tissues or stored, usually as starch. Another direction of phloem conduction is from the root, where sugar is stored as starch, to other parts of the plant, where sugar is used for growth and maintenance of tissues. The vascular cambium, which gives rise to secondary tissues in woody plants, is sandwiched between the xylem and phloem. Because it has an inner core of vascular tissue, the primary eudicot root lacks pith, a ground tissue found in the centers of many stems and roots. Xylem does not form the central tissue in some monocot roots Monocot roots vary considerably in internal structure compared with eudicot roots. The layers found in some monocot roots, starting from the outside, are epidermis, cortex, endodermis, and pericycle (FIG. 36-8). Unlike the xylem in herbaceous eudicot roots, the xylem in many monocot roots does not form a solid cylinder

Stele

Epidermis cell Cortex cell Endodermis cell Pericycle cell

section. As the vascular cambium divides to produce secondary tissues, it eventually forms a cylinder of vascular cambium that continues to divide, producing secondary xylem (wood) to the inside and secondary phloem (inner bark) to the outside. The root increases in girth (thickness), and the vascular cambium continues to move outward. The epidermis, cortex, endodermis, and primary phloem are gradually torn apart as the root increases in girth. The root epidermis is replaced by periderm, composed of cork cells and cork parenchyma, both produced by the cork cambium (the last figure in Table 33-4 shows an LM of periderm). The cork cambium in the root initially arises from regions in the pericycle.

Pith cell Xylem vessel element

Cengage

Phloem cell

250 μm

FIGURE 36-8 LM of a cross section of a monocot root Shown is a greenbrier (Smilax) root. As in herbaceous eudicot roots, the cortex of a monocot root is extensive.

in the center. Instead, the phloem and xylem are in separate, alternating bundles arranged around the central pith, which consists of parenchyma cells. Because virtually no monocots have secondary growth, no vascular cambium exists in monocot roots. Despite their lack of secondary growth, long-lived monocots, such as palms, may have thickened roots produced by a modified form of primary growth in which parenchyma cells in the cortex divide and enlarge.

Woody plants have roots with secondary growth Plants that produce stems with secondary growth also produce roots with secondary growth. These plants—gymnosperms and woody eudicots—have primary growth at apical meristems and secondary growth at lateral meristems. The production of secondary tissues occurs some distance back from the root tips and results from the activity of the same two lateral meristems found in woody stems: the vascular cambium and the cork cambium. Major roots of trees are often massive and have both wood and bark. In temperate climates, the wood of both roots and stems exhibits annual rings in cross section. Before secondary growth starts in a root, the vascular cambium is sandwiched between the primary xylem and the primary phloem (FIG. 36-9). At the onset of secondary growth, the vascular cambium extends out to the pericycle, which develops into vascular cambium opposite the xylem arms. As a result, the pericycle links the separate sections of vascular cambium so that the vascular cambium becomes a continuous, noncircular loop of cells in cross

Some roots are specialized for unusual functions Adventitious roots often arise from the nodes of stems. Many aerial adventitious roots are adapted for functions other than anchorage, absorption, conduction, or storage. Prop roots are adventitious roots that develop from branches or a vertical stem and grow downward into the soil to help support the plant in an upright position (FIG. 36-10a). Prop roots are more common in monocots than in eudicots. Corn and sorghum, both monocots, are herbaceous plants that produce prop roots. Many tropical and subtropical eudicot trees, such as red mangrove, also produce prop roots. The roots of many tropical rainforest trees are shallow and concentrated near the surface in a mat only a few centimeters (an inch or so) thick. The root mat catches and absorbs almost all minerals released from leaves by decomposition. Swollen bases or braces called buttress roots hold the trees upright and aid in the extensive distribution of the shallow roots (FIG. 36-10b). In swampy or tidal environments where the soil is flooded or waterlogged, some roots grow upward until they are above the high-tide level. Even though roots live in the soil, they still require oxygen for aerobic respiration. Flooded soils are depleted of oxygen, so these aerial “breathing” roots, known as pneumatophores, may assist in getting oxygen to the submerged roots (FIG. 36-10c). Pneumatophores, which also help anchor the plant, have a well-developed system of internal air spaces that is continuous with the submerged parts of the root, presumably allowing gas exchange. Black and white mangroves are examples of plants with pneumatophores. Climbing plants and epiphytes, which are plants that grow attached to other plants, have aerial roots that anchor the plant to the bark, branch, or other surface on which they grow. Some epiphytes have aerial roots specialized for functions other than anchorage. Certain epiphytic orchids, for example, have photosynthetic roots (FIG. 36-10d). Epiphytic roots may absorb moisture as well. Some parasitic epiphytes, such as mistletoe, have modified roots that penetrate the host-plant tissues and absorb water. Another plant that starts its life as an epiphyte is the strangler fig, which produces long roots that eventually reach the ground and anchor the plant (now a tree rather than an epiphyte) in the soil. The tree on which the strangler fig originally grew is often killed as the strangler fig grows around it, competing with it for light and other resources and crushing its secondary phloem.

KEY POINT

The production of secondary tissues in roots, like that in stems, is the result of the activity of two lateral meristems—the vascular cambium and the cork cambium.

1

Tissues in primary root.

2

At onset of secondary growth, vascular cambium extends out to pericycle, forming continuous, noncircular loop.

3

Epidermis Cortex Endodermis Pericycle Primary phloem Vascular cambium Primary xylem

Pericycle Primary phloem Vascular cambium Primary xylem

Periderm and remnants of the epidermis, cortex, endodermis, and pericycle

Vascular cambium produces secondary xylem to its inside and secondary phloem to its outside.

Remnant of the primary phloem Secondary phloem Vascular cambium Secondary xylem Primary xylem

4

Periderm and crushed remnants of the primary tissues (outer bark) Secondary phloem (inner bark)

Ring of vascular cambium gradually becomes circular. As vascular cambium continues to divide, epidermis, cortex, and primary phloem located in outer bark are torn apart.

Vascular cambium Secondary xylem (wood) Remnant of the primary xylem

FIGURE 36-9 Development of secondary vascular tissues in a primary root The figures are not drawn to scale because of space limitations; the primary xylem is actually the same size in all four diagrams, but differences in scale make the xylem appear different sizes.

Review ■

■

■ ■ ■

What are the advantages of a taproot system? of a fibrous root system? If you were examining a cross section of a primary root of a flowering plant, how would you determine whether it was a eudicot or a monocot? What is the symplast? the apoplast? How does a herbaceous eudicot root develop secondary tissues? What are the functions of each of the following: prop roots, buttress roots, and pneumatophores?

36.2 ROOT ASSOCIATIONS AND INTERACTIONS ■ ■ LEARNING OBJECTIVE 6 List and describe two mutualistic relationships between roots and other organisms.

As roots of an individual plant extend through the soil, they often encounter the roots of related or unrelated plants. The nature of

Buttress roots

John Arnaldi

Linda R. Berg

Prop roots

(a) Prop roots are adventitious roots that arise near the base of the stem and provide additional support. Screw pine (Pandanus) has an elaborate set of aerial prop roots. Photographed in Kauai, Hawaii.

(b) Tropical rainforest trees typically possess buttress roots that support them in the shallow, often wet soil. Shown are buttress roots on Australian banyan (Ficus macrophylla ). Photographed at Selby Gardens in Sarasota, Florida.

Robert and Linda Mitchell

Aerial roots

John Arnaldi

Pneumatophores

(c) White mangrove (Laguncularia racemosa ) produces pneumatophores, shown protruding from the wet mud in the foreground. Pneumatophores may provide oxygen for roots buried in anaerobic (oxygen-deficient) soil. Photographed in Isla del Carmen, Mexico.

FIGURE 36-10 Specialized roots

(d) The moth orchid (Phalaenopsis hybrid) has photosynthetic aerial roots.

Sheath of fungal hyphae encircles root Fungal hyphae within plant cortical cells

Cabisco / Visuals Unlimited

Robert Knauft / Biology Media / Photo Researchers, Inc.

Fungal hypha between plant cells

250 μm

(a) LM of ectomycorrhizae, fungal associations that form a sheath around the root. The fungal hyphae penetrate the root between cortical cells but do not enter the cells.

100 μm

(b) LM of endomycorrhizae, fungal associations in which the fungal hyphae penetrate root cells. The hyphae aid in delivering and receiving nutrients. Roots of most vascular plant species are colonized by endomycorrhizae.

The roots of terrestrial plant species may form mutualistic— that is, mutually beneficial—relationships with certain soil microorganisms: mycorrhizal fungi and rhizobial bacteria. These mutualistic relationships provide plants with access to essential minerals that are in limited supply in the soil. U.S. Department of Energy

Mycorrhizae facilitate the uptake of essential minerals by roots

(c) Fruiting body of the mycorrhizal fungus Laccaria bicolor, which is colonizing the roots of nearby Douglas fir seedlings.

FIGURE 36-11 Mycorrhizae Mycorrhizae enhance plant growth by providing essential minerals to the roots.

these interactions is largely unknown, but several studies suggest that roots are involved in complex ways, sometimes recognizing the identity of neighboring roots and modifying their growth accordingly. For example, wild strawberry roots have been demonstrated to grow more vigorously when they encounter ground ivy roots; in contrast, root growth of ground ivy is inhibited by strawberry roots. Such root associations affect community dynamics in natural ecosystems and may affect the productivity of agricultural ecosystems.

Subterranean associations between roots and certain fungi, known as mycorrhizae, are found in virtually all terrestrial ecosystems. Mycorrhizae permit the transfer of materials (such as carbon compounds produced by photosynthesis) from roots to the fungus. At the same time, essential minerals, such as phosphorus, move from the fungus to the roots of the host plant. The threadlike body of the fungal partner extends into the soil, extracting minerals well beyond the reach of the plant’s roots. In some mycorrhizae, the fungal mycelium encircles the root like a sheath; in others, the fungus penetrates cell walls of the root cortex and forms branching arbuscules (FIG. 36-11; also see Fig. 29-11). The mycorrhizal relationship is an ancient one: some of the earliest known plant fossils (about 400 million years old) contain mycorrhizal fungi in the roots. The mycorrhizal relationship is mutually beneficial because when mycorrhizae are not present neither the fungus nor the plant grows as well (see Fig. 29-19). The hyphal network of mycorrhizae appears to interconnect the roots of different plant species in the community so that carbon compounds may flow from one plant to another through their mutual fungal partner.

In 2008, an international team of several dozen scientists reported in the journal Nature that the genome of an ectomycorrhizal fungus, Laccaria bicolor, had been sequenced. The scientists identified many genes that code for small secreted proteins that may be involved in establishing the mutualistic relationship between the fungus and its plant partner (see Fig. 36-11c). The functions of these proteins are not known at present, but several are expressed only when the fungus and plant root are in contact to form a mycorrhiza.

Rhizobial bacteria fix nitrogen in the roots of leguminous plants Certain nitrogen-fixing bacteria, collectively called rhizobia, form associations with the roots of leguminous plants—clover, peas, and soybeans, for example. Nodules (swellings) that house millions of the rhizobia develop on the roots (see Fig. 55-9). Evidence indi-

KEY POINT Xylem vessels

cates that the rhizobial relationship evolved much later than the mycorrhizal relationship; unlike mycorrhizae, which infect most plant species, rhizobia almost exclusively infect plants of the legume family, which evolved about 70 million years ago. (A few other plant– bacteria associations that fix nitrogen have also evolved.) As with mycorrhizae, the association between nitrogen-fixing bacteria and the roots of legumes is mutually beneficial. Bacteria receive photosynthetic products from plants while helping plants meet their nitrogen requirements by producing ammonia (NH3) from atmospheric nitrogen. For several decades, biologists have been studying the molecular basis of associations between plants and nitrogen-fixing bacteria. The two partners in this association use cell signaling, that is, a molecular dialogue back and forth between them, to establish contact and develop nodules. When soil contains a low level of nitrogen, legume roots secrete chemical attractants (flavonoids) toward which the rhizobial bacteria swim (FIG. 36-12).

Much of the nitrogen produced by nitrogen-fixing bacteria in legume root nodules is transported in the vascular tissues throughout the plant body, including the seeds. Endodermis

Rhizobial bacteria Cortex

Epidermis Rhizobial bacteria

Phloem cells Pericycle

Root hair 1

Legume root secretes chemical signal that attracts rhizobia.

2

Rhizobia congregate around root hair.

Coiling growth

3

Rhizobia secrete chemicals (nodulation, or nod, factors) that cause root hair to coil around them.

Infection thread

4

Rhizobia enter root hair and move through infection thread (tube surrounded by plasma membrane) that extends down root-hair cell. Rhizobia enclosed in membranes

Developing root nodule

Developing vascular tissues for nodule 5

Rhizobia move from infection thread into cells in cortex; each bacterium is surrounded by a membrane.

6

Chemicals produced by rhizobia induce cell division and formation of root nodule. Some nodule cells are “infected” with numerous bacteria.

FIGURE 36-12 Establishment of a relationship between rhizobial bacteria and leguminous plants

The initial signal molecules produced by rhizobial bacteria are called nodulation factors, or nod factors. These nod factors help root-hair cells in the roots of leguminous plants recognize the presence of the bacteria; recognition is the first step in the infection by rhizobia and the development of root nodules. The nod factor apparently attaches to a plant receptor on the root-hair plasma membrane. This attachment triggers a calcium-dependent signal transduction cascade within root cells that alters gene expression and metabolism, resulting in nodule formation. The plant hormone cytokinin, which triggers cell division (see Chapter 38), is involved in root nodule formation. Some genes are involved in the initiation of both mycorrhizal and rhizobial associations Because both mycorrhizae and rhizobia infect plant roots, some biologists have hypothesized that the genes and signal transduction components involved in mycorrhizal relationships may have been recruited for the rhizobial relationships that evolved much later. There is experimental evidence to support this hypothesis. Some of the same genes that trigger infection and changes in root development to form rhizobial nodules are also involved in establishing mycorrhizal infections. Evidence includes several legume mutants that biologists have identified; these mutants lack the ability to develop nodules for rhizobial bacteria and cannot form mycorrhizal associations. Molecular findings also support the hypothesis.

anchor themselves in soil, and from it they receive water and minerals. Almost all of the elements essential for plant growth are obtained directly from the soil. With few exceptions, plants cannot survive on their own without soil, and because we depend on plants for our food, neither could humans exist without soil. Soils are generally formed from rock (called “parent material”) that is gradually broken down, or fragmented, into smaller and smaller particles by biological, chemical, and physical weathering processes (FIG. 36-13). Two important factors that work together in the weathering of rock are climate and organisms. When plant roots and other organisms living in the soil respire, they produce carbon dioxide (CO2), which diffuses into the soil and reacts with soil water to form carbonic acid (H2CO3). Soil organisms such as lichens produce other kinds of acids. Recall from Chapter 29 that a lichen is a dual organism composed of a fungus and a phototroph (photosynthetic organism). These acids etch tiny cracks, or fissures, in the rock surface; water then seeps into these cracks. If the parent material is located in a temperate climate, the alternate freezing and thawing of the water during winter causes the cracks to enlarge, breaking off small pieces of rock. Small plants can then become established and send their roots into the larger cracks, fracturing the rock further. Topography, a region’s surface features such as the presence or absence of mountains and valleys, is also involved in soil formation. Steep slopes often have little or no soil on them because the soil and rock are continually transported down the slopes by grav-

Review Evaporation and transpiration

What are mycorrhizal fungi? What are rhizobial bacteria? How are mycorrhizae and root nodules similar? How are they different?

■ ■ ■

Precipitation

36.3 THE SOIL ENVIRONMENT ■ ■ LEARNING OBJECTIVES 7 Describe the roles of weathering, organisms, climate, and topography 8 9 10 11

in soil formation. List the four components of soil and give the ecological significance of each. Describe how roots absorb positively charged mineral ions by the process of cation exchange. Soil organisms add Distinguish between macronutrients and organic additions micronutrients. underground. Explain the impacts of mineral depletion and soil Fine particles and erosion on plant growth. solubles are washed downward.

We now examine the soil environment in which most roots live. Soil is a relatively thin layer of Earth’s crust that has been modified by the natural actions of weather, wind, water, and organisms. It is easy to take soil for granted. We walk on and over it throughout our lives but rarely stop to think about how important it is to our survival. Vast numbers and kinds of organisms colonize soil and depend on it for shelter and food. Most plants

Plant litter Earthworms Minerial nutrients

Soil microorganisms

Runoff and erosion are affected by topography.

Water table

Capillary action and evaporation cause some materials to rise.

Parent material Loss of soluble materials in groundwater

FIGURE 36-13 The dynamics of soil formation Weather, climate, topography, and organisms interact with Earth’s crust to form soil, the material that supports plants and many other organisms on land.

ity. Runoff from precipitation tends to amplify erosion on steep slopes. Moderate slopes, in contrast, may encourage the formation of deep soils. The disintegration of solid rock into finer and finer mineral particles in the soil takes an extremely long time, sometimes thousands of years. Soil forms constantly as the weathering of parent material beneath soil that is already formed continues to add new soil.

Clay particle

2+

Ca

2+

Mg K+

K+ Mg

2+

2+

Ca

K+

Ca

Mg

Ca

2+

Mg Mg

H

K+

2+

Mg

2+

Mg

+

2+

Ca

K+

2+

K+

2+

Ca

H

Mg 2+

Ca

+ +

Soil is composed of inorganic minerals, organic matter, air, and water Soil comprises four distinct components—inorganic mineral particles (which make up about 45% of a typical soil), organic matter (about 5%), water (about 25%), and air (about 25%). The plants, animals, fungi, and microorganisms that inhabit soil interact with it, and minerals are continually cycled from the soil to organisms, which use them in their biological processes. When the organisms die, bacteria and other soil organisms decompose the remains, returning the minerals to the soil. The inorganic mineral particles, which come from weathered rock, provide anchorage and essential minerals for plants, as well as pore space for water and air. Because different rocks consist of different minerals, soils vary in mineral composition and chemical properties. Rocks rich in aluminum form acidic soils, for example, whereas rocks that contain silicates of magnesium and iron form soils that may be deficient in calcium, nitrogen, and phosphorus. Also, soils formed from the same kind of parent material may not develop in the same way because other factors, such as weather, topography, and organisms, differ. The texture, or structural characteristic, of a soil is determined by the percentages (by weight) of the different-sized inorganic mineral particles—sand, silt, and clay—in it. The size assignments for sand, silt, and clay give soil scientists a way to classify soil texture. Particles larger than 2 mm in diameter, called gravel or stones, are not considered soil particles because they do not have any direct value to plants. The largest soil particles are called “sand” (0.02 to 2 mm in diameter); the medium-sized particles, “silt” (0.002 to 0.02 mm in diameter); and the smallest particles, “clay” (less than 0.002 mm in diameter). Sand particles are large enough to be seen easily with the eye, and silt particles (about the size of flour particles) can be seen with an ordinary light microscope. Most individual clay particles can be seen only under an electron microscope. A soil’s texture affects many of that soil’s properties, in turn influencing plant growth. The clay component of a soil is particularly important in determining many of its characteristics because clay particles have the greatest surface area of all soil particles. If the surface areas of about 450 g (1 lb) of clay particles were laid out side by side, they would occupy 1 hectare (2.5 acres). Each clay particle has predominantly negative electric charges on its outer surface that attract and reversibly bind cations, which are positively charged mineral ions such as potassium (K+) and magnesium (Mg2+). Because many cations are essential for plant growth, cation adhesion to soil particles is an important aspect of soil fertility. Roots secrete protons (H+), which are exchanged for other positively charged mineral ions adhering to the surface of soil particles, in a process known as cation exchange. These “freed”

2+

Mg

2+

Mg

2+

Ca

2+

2+

Mg

+

H

K+

2+

+

H

K+

Mg

2+

K+

Mg

K+

2+

2+

2+

Mg

2+

Ca K+

2+

H

K+

K+

2+

Ca

2+

2+

Mg

2+

Ca

2+

K+

Root hair

K+

2+

Mg

Ca

2+

Mg

K+

2+

Ca

Ca

2+

Ca

2+

Mg 2+

Ca

K+

2+

Mg

K+

2+

Ca

2+

2+

Mg

K+

Ca

K+

FIGURE 36-14 Cation exchange Negatively charged clay particles bind to positively charged mineral cations, holding them in the soil. Roots pump out protons (H+), which are exchanged for the cations, facilitating their absorption.

ions and the water that forms a film around the soil particles are absorbed by the plant’s roots (FIG. 36-14). In contrast, anions, which are negatively charged mineral ions, are repelled by the negative surface charges of clay particles and tend to remain in solution. Anions such as nitrate (NO3−) are often washed out of the root zone by water moving through the soil. Soil always contains a mixture of different-sized particles, but the proportions vary from one soil to another. A loam, which is an ideal agricultural soil, has an optimum combination of different soil particle sizes: it contains approximately 40% each of sand and silt and about 20% of clay. Generally, the larger particles provide structural support, aeration, and permeability to the soil, whereas the smaller particles bind into aggregates, or clumps, and hold minerals and water. Soils with larger proportions of sand are not as desirable for most plants because they do not hold water and mineral ions well. Plants grown in such soils are more susceptible to drought and mineral deficiencies. Soils with larger proportions of clay are also not desirable for most plants because they provide poor drainage and often do not provide enough oxygen. Clay soils used in agriculture tend to compact, which reduces the number of soil spaces that can be filled by water and air. A soil’s organic matter consists of the wastes and remains of soil organisms The organic matter in soil is composed of litter (dead leaves and branches on the soil’s surface); droppings (animal dung); and the dead remains of plants, animals, and microorganisms in various stages of decomposition. Organic matter is decomposed by microorganisms, particularly bacteria and fungi, that inhabit the soil. During decomposition, essential mineral ions are released into the soil, where they may be bound by soil particles or absorbed by plant roots. Organic matter increases the soil’s water-holding

capacity by acting much like a sponge. For this reason gardeners often add organic matter to soils, especially sandy soils, which are naturally low in organic matter. The partly decayed organic portion of the soil is referred to as humus. Humus, which is not a single chemical compound but a mix of many organic compounds, binds mineral ions and holds water. On average, humus persists in agricultural soil for about 20 years. Certain components of humus may persist in the soil for hundreds of years. Although humus is somewhat resistant to decay, a succession of microorganisms gradually reduces it to carbon dioxide, water, and minerals.

tion of the soil. Some substances completely leach out of the soil because they are so soluble that they migrate down into the groundwater. It is also possible for water that is moving upward through the soil to carry dissolved materials with it.

Soil organisms form a complex ecosystem A single teaspoon of fertile agricultural soil may contain millions of microorganisms, such as bacteria, fungi, algae, protozoa, as well as microscopic nematodes and other worms. Other organisms also colonize the soil ecosystem, including plant roots, earthworms, insects, moles, snakes, and groundhogs (FIG. 36-15). Most numerous in soil are bacteria, which number in the hundreds of millions per gram of soil. Scientists have identified about 170,000 species of soil organisms, but thousands remain to be identified. Little is known about the roles and interactions of soil organisms, in part because it is hard to study their activities under natural conditions. Worms are some of the most important organisms living in soil. Earthworms, probably one of the most familiar soil inhabitants, ingest soil and obtain energy and raw materials by digesting humus. Castings, bits of soil that have passed through the gut of an earthworm, are deposited on or near the soil surface. In this way,

About 50% of soil volume is composed of pore spaces Soil has numerous pore spaces of different sizes around and among the soil particles. Pore spaces occupy roughly 50% of a soil’s volume and are filled with varying proportions of air and water. Both soil air and soil water are necessary to produce a moist but aerated soil that sustains plants and other soil-dwelling organisms. Water is usually held in the smaller pores, whereas air is found in the larger pores. After a prolonged rain, almost all the pore spaces may be filled with water, but water drains rapidly from the larger pore spaces, drawing air from the atmosphere into those spaces. Soil air contains the same gases as atmospheric air, although they are usually present in different proportions. As a result of respiration by soil organisms, there is less oxygen and more carbon dioxide in soil air than in atmospheric air. (Aerobic respiraSurface litter tion uses oxygen and produces carbon dioxide.) Among the important gases in soil are oxygen (O2), required by soil organisms for aerobic respiration; Topsoil nitrogen (N2), used by nitrogen-fixing bacteria; and carbon dioxide (CO2), involved in soil weathering. Soil water originates as precipitaRoot nodules: tion, which drains downward, or as nitrogengroundwater (water stored in porous Subsoil fixing underground rock), which rises upbacteria ward from the water table (the upperBedrock most level of groundwater). Soil water contains low concentrations of dissolved minerals that enter the roots of Mite plants when they absorb water. Water Springtail not bound to soil particles or absorbed Nematode by roots percolates (moves down) Root Bacteria through soil, carrying dissolved minFungus erals with it. The removal of dissolved materials from soil by percolating water is called leaching. The deposition Amoeba of leached material in the lower layers of soil is known as illuviation. Iron and aluminum compounds, humus, FIGURE 36-15 Diversity of life in fertile soil and clay are some illuvial materials Plants, algae, protozoa, fungi, bacteria, earthworms, flatworms, roundworms, insects, spiders and mites, and burrowing animals such as moles and groundhogs live in soil. that can gather in the subsurface por-

minerals from deeper layers are brought to upper layers. Earthworm tunnels serve to aerate the soil, and the worms’ waste products and corpses add organic material to the soil. Ants live in the soil in enormous numbers, constructing tunnels and chambers that help aerate it. Members of soil-dwelling ant colonies forage on the surface for bits of food, which they carry back to their nests. Not all this food is eaten, however, and its eventual decomposition helps increase the organic matter in the soil. Many ants are also indispensable in plant reproduction because they bury seeds in the soil (discussed in Chapter 37).

Soil pH affects soil characteristics and plant growth Acidity is measured using the pH scale, which extends from 0 (extremely acidic) through 7.0 (neutral) to 14.0 (extremely alkaline). The pH of most soils ranges from 4.0 to 8.0, but some soils are outside this range. The soil of the Pygmy Forest in Mendocino County, California, is extremely acidic (pH 2.8 to 3.9). At the other extreme, certain soils in Death Valley, California, have a pH of 10.5. Plants are affected by soil pH, partly because the solubility of certain minerals varies with differences in pH. Soluble minerals can be absorbed by the plant, whereas insoluble forms cannot. At a low pH, for example, aluminum and manganese in soil water are more soluble and are sometimes absorbed by the roots in toxic concentrations. At a higher pH, certain mineral salts essential for plant growth, such as calcium phosphate, become less soluble and thus less available to plants. Soil pH also affects the leaching of minerals. An acidic soil has less ability to bind positively charged ions to it because the soil particles also bind the abundant protons (FIG. 36-16). As a result, certain mineral ions essential for plant growth, such as potassium (K+), are leached more readily from acidic soil. The optimum soil pH for most plant growth is 6.0 to 7.0 because most essential minerals are available to plants in that pH range. Acid precipitation, a type of air pollution in which sulfuric and nitric acids produced by human activities fall to the ground as acid rain, sleet, snow, or fog, can seriously decrease soil pH. Acid precipitation is one of several factors implicated in forest decline, the gradual deterioration, and often death, of trees that has been observed in many European and North American forests in recent decades. Forest decline may be partly the result of soil changes, such as leaching of essential cations, caused by acid precipitation. In central European forests that have experienced forest decline, for example, a strong correlation has been demonstrated between forest damage and soil chemistry altered by acid precipitation.

Soil provides most of the minerals found in plants More than 90 naturally occurring elements exist on Earth, and more than 60 of these, including elements as common as carbon and as rare as gold, have been found in plant tissues. Not all of these elements are essential for plant growth, however.

Nineteen elements have been found essential for most, if not all, plants (TABLE 36-1). Ten of these are required in fairly large quantities (greater than 0.05% dry weight) and are therefore known as macronutrients. Macronutrients include carbon, hydrogen, oxygen, nitrogen, potassium, calcium, magnesium, phosphorus, sulfur, and silicon. The remaining nine micronutrients are needed in trace amounts (less than 0.05% dry weight) for normal plant growth and development. Micronutrients include chlorine, iron, boron, manganese, sodium, zinc, copper, nickel, and molybdenum. Four of the 19 elements—carbon, oxygen, hydrogen, and nitrogen—come directly or indirectly from soil water or from gases in the atmosphere. Carbon is obtained from carbon dioxide in the atmosphere during photosynthesis. Oxygen is obtained from atmospheric oxygen (O2) and water (H2O). Water also supplies hydrogen to the plant. Plants absorb their nitrogen from the soil as ions of nitrogen salts, but the nitrogen in nitrogen salts ultimately comes from atmospheric nitrogen (N2). The remaining 15 essential elements are obtained from the soil as dissolved mineral ions. Their ultimate source is the parent material from which the soil was formed. Let us examine the main functions of essential elements. Carbon, hydrogen, and oxygen are found as part of the structure of all biologically important molecules, including lipids, carbohydrates, nucleic acids, and proteins. Nitrogen is part of proteins, nucleic acids, and chlorophyll. Potassium, which plants use in fairly substantial amounts, is not found in a specific organic compound in plant cells. Instead, it Negatively charged clay particles 2+

Mg

+

H NH4+ + + Na K + 2+ H Ca K+

2+

Mg

NH4+ Ca

2+

(a) In normal soil, positively charged mineral ions are attracted to the negatively charged soil particles.

K+

Na

2+

+Ca

Na

+

H K+ NH +

+

2+

Mg

4

+

H

Al3+

H +

+

H + + H H Al3+

(b) In acidified soil, protons (H+) displace the cations. Aluminum ions released from inorganic mineral particles when the soil becomes acidified also adhere to soil particles.

H

H

+

Al3+

H

+ Al

H

+

H

FIGURE 36-16 How acid alters soil chemistry

+

3+

+

TABLE 36-1

Functions of Elements Required by Most Plants

Element

Taken Up As

Major Functions

Macronutrients Carbon Hydrogen Oxygen Nitrogen Potassium Calcium Magnesium Phosphorus Sulfur Silicon

CO2 H 2O O 2, H 2O NO3−, NH4+ K+ Ca2+ Mg2+ HPO42−, H2PO4− SO42− SiO32−

Component of carbohydrate, lipid, protein, and nucleic acid molecules Component of carbohydrate, lipid, protein, and nucleic acid molecules Component of carbohydrate, lipid, protein, and nucleic acid molecules Component of proteins, nucleic acids, chlorophyll, certain coenzymes Osmotic and ionic balance; opening of stomata; enzyme activator (for 40+ enzymes) In cell walls; involved in membrane permeability; enzyme activator; second messenger in metabolism In chlorophyll; enzyme activator in carbohydrate metabolism (phosphate transfer) In nucleic acids, phospholipids, ATP (energy transfer compound), and coenzymes In certain amino acids and vitamins In cell walls; increased resistance to pests and diseases; important primarily in grasses and sedges

Micronutrients Chlorine Iron Boron Manganese

Cl− Fe2+, Fe3+ H2BO3− Mn2+

Sodium Zinc Copper Nickel Molybdenum

Na+ Zn2+ Cu+, Cu2+ Ni2+ MoO42−

Ionic balance; involved in photosynthesis (production of oxygen) In enzymes and electron transport molecules involved in photosynthesis, respiration, and nitrogen fixation In cell walls; involved in nucleic acid metabolism and in cell elongation In enzymes involved in respiration and nitrogen metabolism; required for photosynthesis (production of oxygen) Involved in photosynthesis (in C4 and CAM plants); substitutes for potassium in osmotic and ionic balance In enzymes involved in respiration and nitrogen metabolism In enzymes involved in photosynthesis In enzymes (urease) involved in nitrogen metabolism; required by nitrogen-fixing bacteria In enzymes involved in nitrogen metabolism

remains as free K+ and plays a key physiological role in maintaining the turgidity of cells because it is osmotically active. The presence of K+ in cytoplasm causes water to pass through the plasma membrane into the cell by osmosis. Potassium is also involved in the opening of stomata. Calcium plays a key structural role as a component of the middle lamella (the cementing layer between cell walls of adjacent plant cells). Calcium ions (Ca2+) are also important in physiological roles in plants, such as altering membrane permeability and as second messengers in various cell signaling responses (see Chapter 6). Magnesium is critical for plants because it is part of the chlorophyll molecule. Phosphorus is a component of nucleic acids, phospholipids (part of cell membranes), and energy transfer molecules such as ATP. Sulfur is essential because it is found in certain amino acids and vitamins. Many plants accumulate silicon in their cell walls and intercellular spaces. Silicon enhances the growth and fertility of some species and may help reinforce cell walls. Chlorine and sodium are micronutrients that help maintain cell turgor. In addition to this osmotic role, chloride (Cl−) and sodium (Na+) ions are essential for photosynthesis. Six of the micronutrients (iron, manganese, zinc, copper, nickel, and molybdenum) are associated with various plant enzymes, often as cofactors, and are involved in certain enzymatic reactions. Boron, present in cell walls, is also involved in nucleic acid metabolism and in cell growth.

How do biologists determine whether an element is essential? It is impossible to conduct mineral nutrition experiments by growing plants in soil because soil is too complex and contains too many elements. Therefore, nutritional studies require special methods. One of the most useful techniques to test whether an element is essential is hydroponics, which is the growing of plants in aerated water to which mineral salts have been added (FIG. 36-17). Hydroponics has commercial applications in addition to its scientific use. Lettuce is one of several crops that is grown hydroponically. If biologists suspect that a particular element is essential for plant growth, they grow plants in a nutrient solution that contains all known essential elements except the one in question. If plants grown in the absence of that element cannot develop normally or complete their life cycle, the element may be essential. Additional criteria are used to confirm whether an element is essential. For example, it must be demonstrated that the element has a direct effect on the plant’s metabolism and that the element is essential for a wide variety of plants.

Soil can be damaged by human mismanagement Soil is a valuable natural resource on which humans depend for food. Many human activities generate or aggravate soil problems, including mineral depletion, soil erosion, and accumulation of salt.

RESEARCH METHOD

Air pump

Using hydroponics, it is possible to grow plants in which only the mineral under investigation is absent from a nutrient solution. (It would be impossible to design such an experiment using soil.) Growing plants hydroponically in the absence of an essential mineral produces recognizable deficiency symptoms.

USDA/Natural Resources Conservation Service

Why Is It Used?

Plant support system Air bubbles

Nutrient solution

How Is It Done?

Air stone

The experimental and control plants are grown in two liquid solutions of mineral nutrients through which air is bubbled to allow the roots to respire. The liquid solution for the experimental plants contains all nutrient minerals known to be essential for plant growth except for one. The liquid solution for the control plants contains all known essential elements, including the one under study.

FIGURE 36-17 Plant nutritional studies in hydroponic growth systems

Mineral depletion may occur in agricultural soils In a natural ecosystem, the essential minerals removed from the soil by plants are returned when the plants or the animals that eat them die and are decomposed. An agricultural system disrupts this pattern of nutrient cycling when the crops are harvested. Plant material containing minerals is removed from the nutrient cycle, and the harvested crops fail to decay and release their nutrients back to the soil. Over time, farmed soil inevitably loses its fertility, that is, its ability to produce abundant crops. Homeowners often mow their lawns and remove the clippings, similarly preventing decomposition and cycling of minerals that were in the grass blades. The essential material (water, sunlight, or some essential element) that is in shortest supply usually limits plant growth. This phenomenon is sometimes called the concept of limiting resources. The three elements that are most often limiting resources for plants are nitrogen, phosphorus, and potassium. To sustain the productivity of agricultural soils, farmers periodically add fertilizers to depleted soils to replace the minerals that limit plant growth.

FIGURE 36-18 Soil erosion in an open field Removal of plant cover exposes bare soil to erosion in heavy rains. Precipitation can cause gullies to enlarge rapidly because they provide channels for the runoff of water.

Soil erosion is the loss of soil from the land Water, wind, ice, and other agents cause soil erosion, the wearing away or removal of soil from the land. Water and wind are particularly effective in moving soil from one place to another. Rainfall loosens soil particles that can then be transported by moving water (FIG. 36-18). Wind loosens soil and blows it away, particularly if the soil is barren and dry. Soil erosion is a natural process that can be greatly accelerated by human activities. Soil erosion is a national and international problem that does not make the headlines very often. The U.S. Department of Agriculture estimates that approximately 4.2 billion metric tons (4.6 billion tons) of topsoil are eroded each year from U.S. croplands and rangelands. Erosion causes a loss in soil fertility because essential minerals and organic matter that are a part of the soil are removed. As a result of these losses, the productivity of eroded agricultural soils declines, and more fertilizer must be used to replace the nutrients lost to erosion. Humans often accelerate soil erosion through poor soil management practices. Agriculture is not the only culprit, because the removal of natural plant communities during surface mining, unsound logging practices (such as clear-cutting large areas of forest), and the construction of roads and buildings also accelerate erosion. Soil erosion has an impact on other natural resources. Sediment that enters streams, rivers, and lakes degrades water quality and fish habitats. Pesticides and fertilizer residues in sediment may add pollutants to the water. Also, when forests are removed within the watershed of a hydroelectric power facility, accelerated soil erosion fills the reservoir behind the dam with sediment much faster than usual. This process reduces electricity production at that facility.

Sufficient plant cover reduces the amount of soil erosion: leaves and stems cushion the impact of rainfall, and roots help hold the soil in place. Although soil erosion is a natural process, abundant plant cover makes it negligible in many natural ecosystems. Salt accumulates in soil that is improperly irrigated Although irrigation improves the agricultural productivity of arid and semiarid lands, it sometimes causes salt to accumulate in the soil, a process called salinization. In a natural scenario, as a result of precipitation runoff, rivers carry dissolved salts away. Irrigation water, however, normally soaks into the soil and does not run off the land into rivers, so when water evaporates, the salt remains behind and accumulates in the soil. Salty soil results in a decline in productivity and, in extreme cases, renders the soil completely unfit for crop production. Most plants cannot obtain all the water they need from salty soil because a water balance problem exists: water moves by osmosis out of plant roots and into the salty soil. Obviously, most plants cannot survive under these conditions (see Fig. 5-14). Plant species that thrive in saline soils have special adaptations that en-

■ ■

able them to tolerate the high amount of salt. Black mangroves, for example, excrete excess salt through their leaves. Most crops, unless they have been genetically selected to tolerate high salt, are not productive in saline soil. Research on the molecular aspects of salt tolerance suggests that mutations in membrane transport proteins confer salt tolerance. Biologists genetically engineered a salt-tolerant variety of Arabidopsis to overexpress a single gene that codes for a sodium transport protein in the vacuolar membrane. These plants can store large quantities of sodium in their vacuoles, thereby tolerating saline soils.

Review What are the four components of soil, and how is each important to plants? What is cation exchange? How do weathering processes convert rock into soil? What are macronutrients and micronutrients? How are minerals lost from the soil?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

36.1 (page 762) 1 Distinguish between taproot and fibrous root systems. ■

A taproot system has one main root (formed from the radicle), from which many lateral roots extend. A fibrous root system has several to many adventitious roots of the same size developing from the end of the stem. Lateral roots branch from these adventitious roots. Learn more about root systems by clicking on the figure in CengageNOW.

2 Compare cross sections of a primary eudicot root and a monocot root, and describe the functions of each tissue. Primary roots have an epidermis, ground tissues (cortex and, in certain roots, pith), and vascular tissues (xylem and phloem). Each root tip is covered by a root cap, a protective layer that covers the delicate root apical meristem and may orient the root so that it grows downward. ■ Epidermis protects the root. Root hairs, short-lived extensions of epidermal cells, aid in absorption of water and dissolved minerals. ■ Cortex consists of parenchyma cells that often store starch. Endodermis, the innermost layer of the cortex, regulates the movement of water and minerals into the root xylem. Cells of the endodermis have a Casparian strip around their radial and transverse walls that is impermeable to water and dissolved minerals. ■

Pericycle, xylem, and phloem collectively make up the root’s stele, or vascular cylinder. Pericycle gives rise to lateral roots and lateral meristems. Xylem conducts water and dissolved minerals; phloem conducts dissolved sugar. ■ Xylem of a herbaceous eudicot root forms a solid core in the center of the root. The center of a monocot root often consists of pith surrounded by a ring of alternating bundles of xylem and phloem. 3 Trace the pathway of water and mineral ions from the soil through the various root tissues, and distinguish between the symplast and apoplast. ■ As water and dissolved mineral ions move from the soil into the root, they pass through the following tissues: root hair/epidermis ¡ cortex ¡ endodermis ¡ pericycle ¡ root xylem. ■ Water and dissolved minerals move through the epidermis and cortex along one of two pathways, the apoplast (along the interconnected porous cell walls) or the symplast (from one cell’s cytoplasm to the next through plasmodesmata). ■

Explore the pathways of water and dissolved minerals in the root by clicking on the figures in CengageNOW.

4 Discuss the structure of roots with secondary growth. ■

Roots of gymnosperms and woody eudicots develop secondary tissues (wood and bark). The production of secondary tissues is the result of the activity of two lateral meristems, the vascular cambium and cork cambium. The vascular cambium produces secondary xylem (wood) and secondary phloem (inner bark). The cork cambium produces periderm (outer bark).

5 Describe at least three roots that are modified to perform uncommon functions. ■ Prop roots develop from branches or from a vertical stem and grow downward into the soil to help support certain plants in an upright position. Buttress roots are swollen bases or braces that support certain tropical rainforest trees that have shallow root systems. Pneumatophores are aerial “breathing” roots that may assist in getting oxygen to submerged roots.

36.2 (page 768) 6 List and describe two mutualistic relationships between roots and other organisms. Mycorrhizae, which are mutually beneficial associations between roots and soil fungi, enhance plant growth by providing essential minerals such as phosphorus to the roots. ■ Root nodules are swellings that develop on roots of leguminous plants and house millions of rhizobia (nitrogen-fixing bacteria). ■

36.3 (page 772) 7 Describe the roles of weathering, organisms, climate, and topography in soil formation. Factors influencing soil formation include parent material (type of rock), climate, organisms, the passage of time, and topography. Most soils are formed from parent material that is broken into smaller and smaller particles by weathering processes. Climate and organisms work together in weathering rock. ■ Soil organisms such as plants, algae, fungi, worms, insects, spiders, and bacteria are important not only in forming soil but also in cycling minerals. ■ Topography, a region’s surface features, affects soil formation. Steep slopes have little or no soil on them, whereas moderate slopes often have deep soils. 8 List the four components of soil and give the ecological significance of each. ■ Soil is composed of inorganic minerals, organic matter, air, and water. Inorganic minerals provide anchorage and essential minerals for plants. Organic matter increases the soil’s water-holding capacity and, as it decomposes, releases essential minerals into the soil. Soil ■

air provides oxygen for soil organisms to use during aerobic respiration. Soil water provides water and dissolved minerals to plants and other organisms. 9 Describe how roots absorb positively charged mineral ions by the process of cation exchange. ■ Cations, positively charged mineral ions, are attracted and reversibly bound to clay particles, which have predominantly negative charges on their outer surfaces. In cation exchange, roots secrete protons (H+), which are exchanged for other positively charged mineral ions, freeing them into the soil water to be absorbed by roots. 10 Distinguish between macronutrients and micronutrients. ■ Plants require 19 essential elements for normal growth. Ten elements are macronutrients: carbon, hydrogen, oxygen, nitrogen, potassium, calcium, magnesium, phosphorus, sulfur, and silicon. Macronutrients are required in fairly large quantities. ■ Nine elements are micronutrients: chlorine, iron, boron, manganese, sodium, zinc, copper, nickel, and molybdenum. Micronutrients are needed in trace amounts. 11 Explain the impacts of mineral depletion and soil erosion on plant growth. ■ Mineral depletion may occur in soils that are farmed because the natural pattern of nutrient cycling is disrupted when crops are harvested (and not allowed to decompose into the soil). ■ Soil erosion is the removal of soil from the land by agents such as water and wind. Erosion causes a loss in soil fertility because essential minerals and organic matter that are a part of the soil are removed.

T E S T YO U R U N D E R S TA N D I N G 1. One main root, formed from the enlarging embryonic root, with many smaller lateral roots branching off it is (a) a fibrous root system (b) an adventitious root system (c) a taproot system (d) a storage root system (e) a prop root system 2. Root hairs (a) cover and protect the delicate root apical meristem (b) increase the absorptive capacity of roots (c) secrete a waxy cuticle (d) orient the root so it grows downward (e) store excess sugars produced in the leaves 3. Certain plants adapted to flooded soil produce aerial “breathing” roots known as (a) fibrous roots (b) pneumatophores (c) mycorrhizae (d) nodules (e) prop roots

6. Plants obtain positively charged mineral ions from clay particles in the soil by cation exchange, in which (a) roots passively absorb the positively charged mineral ions (b) mineral ions flow freely along porous cell walls (c) roots secrete protons, which free other positively charged mineral ions to be absorbed by roots (d) the Casparian strip effectively blocks the passage of water and mineral ions along the endodermal cell wall (e) a well-developed system of internal air spaces in the root allows both gas exchange and cation exchange 7. The cell layer from which lateral roots originate is the (a) epidermis (b) cortex (c) endodermis (d) pericycle (e) vascular cambium

4. The waterproof region around the radial and transverse walls of endodermal cells is the (a) Casparian strip (b) pericycle (c) apoplast (d) symplast (e) pneumatophore

8. Mutually beneficial associations between certain soil fungi and the roots of most plant species are called (a) mycorrhizae (b) pneumatophores (c) nodules (d) rhizobia (e) humus

5. The apoplast is (a) a layer of cells that surround the vascular region in roots (b) the layer of cells just inside the endodermis (c) a system of interconnected plant cell walls through which water moves (d) the central cylinder of the root that comprises the vascular tissues (e) a continuum of cytoplasm of many cells, all connected by plasmodesmata

9. Which of the following statements about soil is true? (a) pore spaces are always filled with about 50% air and 50% water (b) a single teaspoon of fertile agricultural soil may contain up to several hundred living microorganisms (c) the texture of a soil is determined by the soil’s pH (d) a soil’s organic matter includes litter, droppings, and the dead remains of plants, animals,

and microorganisms (e) soil formation is unaffected by a region’s climate or topography 10. Carbon, hydrogen, oxygen, nitrogen, potassium, calcium, magnesium, phosphorus, sulfur, and silicon are collectively known as (a) micronutrients (b) microvilli (c) micronuclei (d) macronuclei (e) macronutrients 11. Corn, sorghum, and red mangrove are plants that have (a) prop roots (b) buttress roots (c) pneumatophores (d) storage roots (e) root nodules 12. Label the various tissues of this herbaceous eudicot root. Use Figure 36-3 to check your answers.

CRITICAL THINKING

2. How would you distinguish between a root hair and a small lateral root? 3. You are given a plant part that was found growing in the soil and are asked to determine whether it is a root or an underground stem. How would you identify the structure without a microscope? with a microscope? 4. How would you design an experiment to determine whether gold is essential for plant growth? What would you use for an experimental control? 5. Why does overwatering a plant often kill it? 6. Explain why, once secondary growth has occurred, that portion of the root is no longer involved in absorption. Where does absorption of water and dissolved minerals occur in plants that have roots with secondary growth? 7. EVOLUTION LINK. A barrel cactus that is 60 cm tall and 30 cm in diameter has roots more than 3 m long. However, all the plant’s roots are found in the soil at a depth of 5 to 15 cm. What possible adaptive value does such a shallow root system confer on a desert plant? 8. ANALYZING DATA. When soil has become polluted with hazardous materials such as heavy metals, it is sometimes costeffective to construct an artificial marsh around the site. Water is pumped through the contaminated soil, picking up the hazard-

ous material, and then the water is passed through the artificial wetland. In one experiment, scientists compared the effectiveness of the roots of two different marsh plants (cattails and common reeds) in absorbing the heavy metal selenium. Which plant’s roots removed the most selenium? What should be done to the plants after they have absorbed selenium? 100 Control Selenium concentration in water passing through artificial marsh (μg/L)

1. A mesquite root is found penetrating a mine shaft about 46 m (150 ft) below the surface of the soil. How could you determine when the root first grew into the shaft? (Hint: Mesquite is a woody plant.)

80

Cattails (Typha latifolia)

60

40

20

0

Common reeds (Phragmites australis) Inlet

1

2

3

4

Outlet

Sample locations

Additional questions are available in CengageNOW at www.cengage.com/ login.

Reproduction in Flowering Plants

Dr. Jeremy Burgess/Science Photo Library/Photo Researchers, Inc.

37

KEY CONCEPTS

Flower of the common chickweed. This plant (Stellaria media) is a weedy annual native to Europe but widespread in North America. Each flower has five green sepals, five deeply notched yellow petals, three to five pollen-bearing stamens (this flower has three), and a single pistil.

F

lowering plants, or angiosperms, include about 300,000 species and are the largest, most successful group of plants. You may have

37.1 The flower is the site of sexual reproduction in an-

admired flowers for their fragrances as well as their appealing colors

giosperms. A typical flower consists of four whorls: sepals, petals, stamens, and carpels.

and varied shapes (see photograph). The biological function of flowers

37.2 Pollen grains are transported to stigmas by a variety

likelihood that pollen grains, which produce sperm cells, will be carried

of agents, such as animals and wind.

37.3 Double fertilization results in a plant embryo and endosperm. The seed is a mature ovule, and the fruit is a mature ovary.

37.4 Following a period of dormancy, a seed germinates,

is sexual reproduction. Their colors, shapes, and fragrances increase the from one plant to another. Sexual reproduction in plants includes meiosis and the fusion of reproductive cells—egg and sperm cells, collectively called gametes. The union of gametes, called fertilization, occurs within the flower’s ovary. Sexual reproduction offers the advantage of new gene combinations,

or sprouts. The young seedling becomes anchored in the ground and begins to photosynthesize.

not found in either parent, that may make an individual plant better suited

37.5 Many vegetative organs (roots, stems, and leaves)

to its environment. These new combinations result from the crossing-over

are modified for asexual reproduction.

and independent assortment of chromosomes that occur during meiosis,

37.6 Sexual reproduction in flowers results in genetic vari-

before the production of egg and sperm cells (see Chapter 11).

ability in the offspring, whereas asexual reproduction in vegetative structures generally results in individuals that are genetically identical.

Many flowering plants also reproduce asexually. Asexual reproduction often does not involve the formation of flowers, seeds, and fruits. Instead, offspring generally form when a vegetative organ (such as a stem, root, or leaf) expands, grows, and then becomes separated from

the rest of the plant, often by the death of tissues. Because asexual reproduction requires only one parent and no meiosis or fusion of gametes occurs, the offspring of asexual reproduction are virtually genetically identical to one another and to the parent plant.1 This chapter examines both sexual and asexual reproduction in flowering plants, including floral adaptations that are important in pollination; seed and fruit structure and dispersal; germination and early growth; and several kinds of asexual reproduction. We conclude with a discussion of the evolutionary advantages and disadvantages of sexual and asexual reproduction.

37.1 THE FLOWERING PLANT LIFE CYCLE ■ ■ LEARNING OBJECTIVES 1 2

Describe the functions of each part of a flower. Identify where eggs and pollen grains are formed within the flower.

In Chapters 27 and 28 you learned that angiosperms and other plants undergo a cyclic alternation of generations in which they spend a portion of their life cycle in a multicellular haploid stage and a portion in a multicellular diploid stage. The haploid portion, called the gametophyte generation, gives rise to gametes by mitosis. When two gametes fuse during fertilization, the diploid portion of the life cycle, called the sporophyte generation, begins. The sporophyte generation produces haploid spores by meiosis. Each spore has the potential to give rise to a gametophyte plant, and the cycle continues. In flowering plants the diploid sporophyte generation is larger and nutritionally independent. The haploid gametophyte generation, which is located in the flower, is microscopic and nutritionally dependent on the sporophyte. We say more about alternation of generations in flowering plants after our discussion of the role of flowers as reproductive structures. (It may be helpful for you to review Figure 28-13, which shows the main stages in the flowering plant life cycle.)

Flowers develop at apical meristems How does a plant “know” it is time to start forming flowers? Correct timing in the switch from vegetative to reproductive development is crucial to ensure reproductive success. A variety of environmental cues, such as temperature and day length, ensure proper timing, and different species are adapted to respond to distinct environmental cues. These environmental signals interact with a variety of plant hormones and developmental pathways. In recent years some biologists have focused on the molecular aspects of developmental pathways that initiate flowering in plants such as the model organism Arabidopsis. When environ1

Somatic mutations can result in some variability among asexually derived offspring.

mental conditions induce flowering, many genes are activated or inactivated. One gene, the Flowering Locus C (FLC) gene, codes for a transcription factor that represses flowering. Another gene, called Flowering Locus D (FLD) gene, codes for a protein that removes acetyl groups from histones in the chromatin where the FLC gene is located. When deacetylation occurs, the FLC gene is not transcribed (that is, the repressive transcription factor is not produced), and the shoot apical meristem undergoes a transition from vegetative growth to reproductive growth. It is intriguing that the plant FLD protein is homologous to a mammalian protein that also removes acetyl groups from chromatin. Other genes are also involved in the initiation of flowering, and this area remains a focus of active research interest. In Chapter 38 we discuss the initiation of flowering further.

Each part of a flower has a specific function Flowers are reproductive shoots, usually consisting of four kinds of organs—sepals, petals, stamens, and carpels—arranged in whorls (circles) on the end of a flower stalk (FIG. 37-1; also see Fig. 28-11). In flowers with all four organs, the normal order of whorls from the flower’s periphery to the center (or from the flower’s base upward) is as follows: sepals ¡ petals ¡ stamens ¡ carpels

The tip of the stalk enlarges to form a receptacle on which some or all of the flower parts are borne. All four floral parts are important in the reproductive process, but only the stamens (the “male” organs) and carpels (the “female” organs) participate directly in sexual reproduction—sepals and petals are sterile. Sepals, which constitute the outermost and lowest whorl on a floral shoot, cover and protect the flower parts when the flower is a bud. Sepals are leaflike in shape and form and are often green. Some sepals, such as those in lily flowers, are colored and resemble petals (FIG. 37-2). The collective term for all the sepals of a flower is calyx. The whorl just inside and above the sepals consists of petals, which are broad, flat, and thin (like sepals and leaves) but widely varied in shape and frequently brightly colored, which attracts pollinators. Petals play an important role in ensuring that sexual reproduction will occur. Sometimes petals fuse to form a tube or other floral shape. The collective term for all the petals of a flower is corolla. Just inside and above the petals are the stamens, the male reproductive organs. Each stamen has a thin stalk, called a filament, at the top of which is an anther, a saclike structure in which pollen grains form. For sexual reproduction to occur, pollen grains must be transferred from the anther to the female reproductive structure (the carpel), usually of another flower of the same species. At first, each pollen grain consists of two cells surrounded by a tough outer wall. One cell, the generative cell, divides mitotically to form two nonflagellate male gametes, known as sperm cells. The other cell, the tube cell, produces a pollen tube, through which the sperm cells travel to reach the ovule. One or more carpels, the female reproductive organs, are located in the center or top of most flowers. Carpels bear ovules,

which are structures with the potential to develop into seeds. The carpels of a flower may be separate or fused into a single structure. The female part of the flower, often called a pistil, may be a single carpel (a simple pistil) or a group of fused carpels (a compound pistil) (see Fig. 28-12). Each pistil has three sections: a stigma, on which the pollen grains land; a style, a necklike structure through

which the pollen tube grows; and an ovary, a juglike structure that contains one or more ovules and can develop into a fruit. Considerable variation in flower structure occurs from one species to another, but flower structure is virtually constant within a given species. For that reason, flower structure is important in plant identification and, if flowers are present, flowering plant species are easy to evaluate in the field.

Darwin Dale / Photo Researchers, Inc.

Female gametophytes are produced in the ovary, male gametophytes in the anther Before we proceed, it may be helpful to relate the stages in alternation of generations to floral structure. As discussed in Chapters 27 and 28, angiosperms and certain other plants are heterosporous and produce two kinds of spores: megaspores and microspores (FIG. 37-3). Each young ovule within an ovary contains a diploid cell, the megasporocyte, which undergoes meiosis to produce four haploid megaspores. Three of these usually disintegrate, and the fourth, the functional megaspore, divides mitotically to produce a multicellular female gametophyte, also called an embryo sac. The female gametophyte, which is embedded in the ovule, typically contains seven cells with eight haploid nuclei. Six of these cells, including the egg cell (the female gamete), contain a single nucleus each; a large central cell has two nuclei, called polar nuclei. The egg and both polar nuclei participate directly in fertilization. (a) An Arabidopsis thaliana flower. Female floral parts

Male floral parts Pollen grain (each will produce two sperm cells)

Stigma Style PISTIL (consisting of one or more carpels)

Anther STAMEN Filament

Ovary Ovules (each producing one egg cell)

Petal

Receptacle

Sepal

Peduncle

(b) Cutaway view of an Arabidopsis flower. Each flower has four sepals (two are shown), four petals (two are shown), six stamens, and one long pistil. Four of the stamens are long, and two are short (two long and two short are shown). Pollen grains develop within sacs in the anthers. In Arabidopsis, the compound pistil consists of two carpels that each contain numerous ovules.

FIGURE 37-1 Animated Floral structure

© Holmes Garden Photos/ Alamy

Many plants have mechanisms that prevent self-pollination

FIGURE 37-2 Lily flowers In lily (Lilium) the three sepals and three petals are similar in size and color.

Pollen sacs within the anther contain numerous diploid cells called microsporocytes, each of which undergoes meiosis to produce four haploid cells called microspores. Each microspore divides mitotically to produce an immature male gametophyte, also called a pollen grain, that consists of two cells, the tube cell and the generative cell. The pollen grain becomes mature when its generative cell divides to form two nonmotile sperm cells.

Review ■ ■ ■ ■

How do petals differ from sepals? How are they similar? How do stamens differ from carpels? How are they similar? What are female gametophytes, and where are they formed? What are male gametophytes, and where are they formed?

37.2 POLLINATION ■ ■ LEARNING OBJECTIVES 3

Compare the evolutionary adaptations that characterize flowers pollinated in different ways (by insects, birds, and bats).

4 Define coevolution and give examples of ways that plants and their animal pollinators have affected one another’s evolution.

Before fertilization can occur, pollen grains must travel from the anther (where they form) to the stigma. The transfer of pollen grains from anther to stigma is known as pollination. Plants are self-pollinated if pollination occurs within the same flower or a different flower on the same individual plant. When pollen grains are transferred to a flower on another individual of the same species, the plant is cross-pollinated. Flowering plants accomplish pollination in a variety of ways. Beetles, bees, flies, butterflies, moths, wasps, and other insects pollinate many flowers. Animals such as birds, bats, snails, and small nonflying mammals (rodents, primates, and marsupials) also pollinate plants. Wind is an agent of pollination for certain flowers; and water, for a few aquatic flowers.

In plant sexual reproduction, the two gametes that unite to form a zygote may be from the same parent or from two different parents. The combination of gametes from two different parents increases the variation in offspring, and this variation may confer a selective advantage. Some offspring, for example, may be able to survive environmental changes better than either parent can. Evolution has resulted in a variety of mechanisms that prevent self-pollination and thus prevent inbreeding, which is the mating of genetically similar individuals. Inbreeding can increase the concentration of harmful genes in the offspring. Some plants, such as asparagus and willow, have separate male and female individuals; the male plants have staminate flowers that lack carpels, and the female plants have pistillate flowers that lack stamens. Other species have flowers with both stamens and pistils, but the pollen is shed from a given flower either before or after the time when the stigma of that flower is receptive to pollen. These characteristics promote outcrossing (also called outbreeding), which is the mating of dissimilar individuals. Many species have genes for self-incompatibility, a genetic condition in which the pollen is ineffective in fertilizing the same flower or other flowers on the same plant. In other words, an individual plant can identify and reject its own pollen. Genes for self-incompatibility usually inhibit the growth of the pollen tube in the stigma and style, thereby preventing delivery of sperm cells to the ovules. Self-incompatibility, which is more common in wild species than in cultivated plants, ensures that reproduction occurs only if the pollen comes from a genetically different individual. In plants such as oilseed rape, self-incompatibility is based on a high degree of variation at a particular locus called the S locus; the many alleles at this locus are designated S1, S2 , S3 , S4 , and so on. Here is an example of how the S locus blocks self-fertilization: A plant with the genotype S1S2 produces pollen grains that land on a stigma of another plant with the genotype S1S3. In this case, the presence of the S1 allele in both the pollen and stigma triggers a self-recognition signaling cascade in surface cells of the stigma. As a result, the stigma cells do not undergo changes that allow the pollen grain to grow a pollen tube. Therefore, fertilization does not occur. The molecular basis of self-incompatibility in Arabidopsis and related plants is an area of active scientific interest. Arabidopsis can self-pollinate. Biologists have determined that the genes involved in self-incompatibility and outcrossing exist in Arabidopsis but have undergone mutations so that they are no longer functional. Thus, it appears that self-incompatibility in these plants is the ancestral (normal) condition.

Flowering plants and their animal pollinators have coevolved Animal pollinators and the plants they pollinate have had such close, interdependent relationships over time that they have affected the evolution of certain physical and behavioral features in one another. The term coevolution describes such reciprocal adaptation, in which two species interact so closely that they become increasingly adapted to each other as they each undergo evo-

KEY POINT

In alternation of generations in flowering plants, the female and male gametophytes are microscopic and nutritionally dependent on the sporophyte plant.

Young ovule (megasporangium) Megasporocyte

Young pollen sac with numerous microsporocytes

Meiosis Microsporocyte Functional megaspore

Meiosis

Disintegrating megaspores (3)

Microspores (4)

Mitosis (3 divisions)

Single microspore

Antipodal cells

Mitosis Female gametophyte (embryo sac)

Polar nuclei in a central cell Synergids Integuments

Egg

Pollen grain (immature male gametophyte)

Generative cell (will divide to form 2 sperm cells) Tube cell

FIGURE 37-3 Animated Development of female and male gametophytes (Left side) The female gametophyte, or embryo sac, develops within the ovule. (Right side) The immature male gametophytes, or pollen grains, develop within pollen sacs in the anthers.

lutionary change by natural selection. We now examine some of the features of flowers and their pollinators that may be the products of coevolution. Flowers pollinated by animals have various features to attract their pollinators, including showy petals (a visual attractant) and

scent (an olfactory attractant) (FIG. 37-4 and TABLE 37-1). One reward for the animal pollinator is food. Pollen grains are a proteinrich food for many animals. As they move from flower to flower searching for food, pollinators inadvertently carry along pollen grains on their body parts, helping the plants reproduce sexually.

Dr. John Brackenbury/ Science Photo Library/Photo Researchers, Inc.

Ed Reschke / Peter Arnold, Inc.

(b) An eastern tiger swallowtail butterfly (Pterourus glaucas) obtains nectar from butterfly weed (Asclepias tuberosa) flowers. Photographed in Michigan.

Linda R. Berg

© Graphic Science / Alamy

(a) A honeybee (Apis mellifera ) pollinates a Scotch broom flower (Cytisus scoparius ). When the bee presses in to obtain nectar, the stamens spring out, dropping pollen grains on the bee’s body.

(e) Ultraviolet markings on the evening primrose (Oenothera sp.) draw attention to the center of the flower, where insects find nectar and pollen grains. The human eye perceives the flower as solid yellow (left). The same flower viewed under ultraviolet radiation (right) provides clues about how the insect eye perceives it. The light blue portions of the petals may appear purple to a bee, whereas the dark blue inner parts may appear yellow.

FIGURE 37-4 Flowers and their animal pollinators

(d) The carrion flower (Stapelia sp.), which has petals that resemble dried blood, emits a putrid scent. This desert plant is pollinated by flies. Photographed in the Fairchild Tropical Garden in Miami, Florida.

© Merlin Tuttle /Bat Conservation International /Photo Researchers, Inc.

Thomas Eisner, Cornell University

Thomas Eisner, Cornell University

(c) An Australian long-nosed lycid beetle (Metriorrhynchus rhipidus) obtains nectar from an Australian tea-tree (Leptospermum laevigatum).

(f) A lesser longnosed bat (Leptonycteris curasoae) obtains nectar from a cardon cactus flower (Pachycereus ) and transfers pollen as it moves from flower to flower. Bats pollinate several hundred species of plants.

(h) An orchid flower (Ophrys scolopax) that mimics female bees attracts male bees, thereby accomplishing pollination.

FIGURE 37-4 (continued)

TABLE 37-1

Nuridsany et Perrenou / Photo Researchers, Inc.

© Malcolm Schuyl / Alamy

(g) A broad-billed hummingbird obtains nectar from a desert flower, ocotillo (Fouquieria splendens), in Arizona. The pollen grains on the bird’s feathers are carried to the next plant.

Floral Characteristics Associated with Various Animal Pollinators

Animal Pollinator

Common Flower Color

Scent

Pollinator Reward

Time of Flowering

Bee Butterfly Moth Carrion fly Beetle Bird Bat

Yellow, blue, purple Red, pink White Reddish-brown, purple Green, white Red, pink White

Strong, floral Weak, floral Strong, sweet Strong, rotten Strong, various scents No scent Strong, musklike or sweet

Nectar, pollen Nectar Nectar No reward Nectar, pollen Nectar Nectar, pollen

Day Day Dusk, night Day, night Day, night Day Night

Some flowers produce nectar, a sugary solution, in special floral glands called nectaries. Pollinators use nectar, which is also rich in amino acids, alkaloids, and other floral chemicals, as an energyrich food. Some nectars contain alkaloids such as nicotine to prevent animal pollinators from drinking all the nectar in a single visit. (Nicotine is poisonous.) Thus, pollinators are encouraged to visit multiple flowers, and each flower has multiple pollinator visitors. Biologists estimate that insects pollinate about 70% of all flowering plant species. Bees are particularly important as pollinators of crop plants. Crops pollinated by bees provide about 30% of human food. Plants pollinated by insects often have blue or yellow petals. The insect eye does not see color the same way the human eye does. Most insects see well in the violet, blue, and yellow range of visible light but do not perceive red as a distinct color. So flowers pollinated by insects are not usually red. Insects also see in the ultraviolet range, wavelengths that are invisible to the human eye. Insects see ultraviolet radiation as a color called bee’s purple. Many

flowers have dramatic UV markings that may or may not be visible to humans but that direct insects to the center of the flower where the pollen grains and nectar are. Insects have a well-developed sense of smell, and many insectpollinated flowers have a strong scent that may be pleasant or foul to humans. The carrion plant, for example, is pollinated by flies and smells like the rotting flesh in which flies lay their eggs. As flies move from one reeking flower to another looking for a place to lay their eggs, they transfer pollen grains. Birds such as hummingbirds are important pollinators. Flowers pollinated by birds are usually red, orange, or yellow because birds see well in this range of visible light. Because most birds do not have a strong sense of smell, bird-pollinated flowers usually lack a scent. Bats, which feed at night and do not see well, are important pollinators, particularly in the tropics. Bat-pollinated flowers bloom at night and have dull white petals and a strong nighttime output of

about 25% of lobelia species have become extinct. The ‘i‘iwi now feeds largely on the flowers of the ‘ohi‘a tree, which lacks petals, and the ‘i‘iwi bill appears to be adapting to this change in feeding preference. Comparison of the bills of ‘i‘iwi museum specimens collected in 1902 with the bills of live birds captured in the 1990s showed that ‘i‘iwi bills were about 3% shorter in the 1990s than in 1902. Plants and the behavior of their animal pollinators have coevolved, sometimes in bizarre and complex ways. The flowers of certain orchids (Ophrys) resemble female bees in coloring and shape. The unpollinated flowers secrete a scent similar to that produced by female bees, and the males are irresistibly attracted to it. The resemblance between Ophrys flowers and female bees is so strong that male bees mount the flowers and try to copulate with them. During this pseudocopulation, a pollen sac usually attaches to the back of the bee. When the frustrated bee departs and tries to copulate with another orchid flower, pollen grains are transferred to that flower. Interestingly, once a flower has been pollinated, it emits a scent like that released by female bees that have already mated. Male bees have no interest in visiting flowers that have been pollinated (just as they lose interest in female bees that have already been inseminated).

scent that usually smells like fermented fruit. Nectar-feeding bats are attracted to the flowers by the scent; they lap up the nectar with their long, extendible tongues. As they move from flower to flower, they transfer pollen grains. An unusual adaptation that encourages pollination by bats has evolved in the tropical vine Mucana holtonii. When the pollen in a given flower is mature, a concave petal lifts up. The petal bounces the echo from the bat’s echo-locating calls back to the bat, helping it find the flower. Specialized features such as petals, scent, and nectar to attract pollinators have coevolved with adaptations of animal pollinators. Specialized body parts and behaviors adapted animals to aid pollination as they obtain nectar and pollen grains as a reward. For example, coevolution has selected for bumblebees’ hairy bodies, which catch and hold the sticky pollen grains for transport from one flower to another. Coevolution may have led to the long, curved beaks of the ‘i‘iwi, one of the Hawaiian honeycreepers, that inserts its beak into tubular flowers to obtain nectar (see Figs. 1-13b and 20-18). The long, tubular corolla of the flowers that ‘i‘iwis visit probably also came about through coevolution. During the 20th century, some of the tubular flower species (such as lobelias) became rare, largely as a result of grazing by nonnative cows and feral goats, and

KEY EXPERIMENT QUESTION: Can a single-gene mutation in flower color affect animal pollinators? HYPOTHESIS: If flower color is changed by a mutation, pollinator preferences for the resulting flower may change.

Wild-type M. lewisii

Wild-type M. cardinalis

Altered M. lewisii

Nature, Vol. 426, November 13, 2003

Nature, Vol. 426, November 13, 2003

Nature, Vol. 426, November 13, 2003

Nature, Vol. 426, November 13, 2003

EXPERIMENT: Mimulus lewisii has violet-pink flowers and is pollinated by bumblebees; the closely related M. cardinalis has orange-red flowers and is pollinated by hummingbirds. Alleles at the same locus in both species affect flower color. Researchers transferred the allele for orange-red flowers from M. cardinalis to M. lewisii, which then produced yellow-orange petals. They also transferred the allele for pink flowers from M. lewisii to M. cardinalis, which then produced dark pink petals.

Altered M. cardinalis

RESULTS AND CONCLUSION: The modified M. lewisii flowers were yellow-orange in color and had 68 times as many hummingbird visits as the unaltered M. lewisii flowers. The modified M. cardinalis flowers had dark pink petals and were visited by 74 times as many bumblebees as the unaltered M. cardinalis flowers. Thus, a single mutation in a gene for flower color resulted in a shift in animal pollinators.

FIGURE 37-5 Studying how pollinator preferences may have evolved Biologists from the University of Washington and Michigan State University used monkey flowers to determine if a single gene mutation would have any effect on animal pollinators.

stigmas, presumably to trap wind-borne pollen grains. Because wind pollination is a hit-or-miss affair, the likelihood of a particular pollen grain landing on a stigma of the same species of flower is slim. Wind-pollinated plants produce large quantities of pollen grains, which increase the likelihood that some pollen grains will land on the appropriate stigma.

A single mutation in a gene for flower color can result in a shift in animal pollinators Biologists used two species of monkey flowers, both native to western North America, to determine if a single gene mutation would have any effect on animal pollinators. The wild-type Mimulus lewisii has violet-pink flowers and is pollinated by bumblebees, whereas the closely related M. cardinalis has orange-red flowers and is pollinated by hummingbirds. Alleles at the same locus in both species affect flower color. When the researchers transferred the allele for orange-red flowers from M. cardinalis to M. lewisii, the resulting M. lewisii flowers were yellow-orange instead of pink. These flowers were visited by 68 times as many hummingbirds as the wild-type M. lewisii flowers (FIG. 37-5). A shift in pollinators also occurred when M. cardinalis plants received the allele for pink flowers from M. lewisii. These M. cardinalis flowers were a dark pink in color and had 74 times as many bumblebee visits as the wild-type M. cardinalis flowers. Biologists think that hummingbird-pollinated flowers arose from insect-pollinated flowers many times during the course of flowering plant evolution. These data suggest that a single gene mutation affecting petal color could have been partly responsible for the pollinator shift that resulted in the evolution of two species, M. lewisii and M. cardinalis, from their insect-pollinated common ancestor.

Review ■

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37.3 FERTILIZATION AND SEED/ FRUIT DEVELOPMENT ■ ■ LEARNING OBJECTIVES 5 Distinguish between pollination and fertilization. 6 Trace the stages of embryo development in flowering plants, and list and define the main parts of seeds.

7

Some flowering plants depend on wind to disperse pollen

8

Dr. Jeremy Burgess/Science Photo Library/Photo Researchers, Inc.

Some flowering plants, such as grasses, ragweed, maples, and oaks, are pollinated by wind. Wind-pollinated plants produce many small, inconspicuous flowers (FIG. 37-6). They do not produce large, colorful petals, scent, or nectar. Some have large, feathery

FIGURE 37-6 Wind pollination Each cluster of male oak flowers (Quercus) dangles from a tree branch and sheds a shower of pollen when the wind blows. These flowers lack petals.

A flower has yellow petals, a pleasant scent, and sugary nectar. How is it probably pollinated? A flower has small, inconspicuous flowers without petals, a scent, or nectar. How is it probably pollinated? What is coevolution?

Explain the relationships among the following: ovules, ovaries, seeds, and fruits. Distinguish among simple, aggregate, multiple, and accessory fruits.

After pollen grains have been transferred from anther to stigma, the tube cell, one of the two cells in the pollen grain, grows a thin pollen tube down through the style and into an ovule in the ovary. How does the pollen tube “know” where to grow? Scientists have found that molecular signals from the female gametophyte guide the growing pollen tube toward the ovule. In 2001 the synergid cells located adjacent to the egg were identified as the source of these molecular signals. The chemical nature of these pollen tube attractants was not known until 2009, when two attracting signals (called LUREs), both small polypeptides, were identified in wishbone flowers (Torenia fournieri) and reported in the journal Science. Once a pollen tube penetrates the ovule, the attracting signals are no longer produced. As a result, only one pollen tube enters each female gametophyte. The second cell within the pollen grain divides to form two male gametes (the sperm cells), which move down the pollen tube and enter the ovule (FIG. 37-7).

A unique double fertilization process occurs in flowering plants The egg within the ovule unites with one of the sperm cells, forming a zygote (fertilized egg) that develops into an embryonic plant contained in the future seed. The two polar nuclei in the central cell of the ovule fuse with the second sperm cell to form the first cell of the triploid (3n) endosperm, the tissue with nutritive and hormonal functions that surrounds the developing embryonic plant in the seed. This process, in which two separate cell fusions occur, is called double fertilization. It is, with few exceptions, unique to flowering plants. (A type of double fertilization has been reported in two gymnosperm species, Ephedra nevadensis and Gnetum gnemon.)

After double fertilization, the ovule develops into a seed and the surrounding ovary develops into a fruit (see discussion in Chapter 28).

Embryonic development in seeds is orderly and predictable Flowering plants package a young plant embryo, complete with stored nutrients, in a compact seed, which develops from the ovule after fertilization. The nutrients in seeds are not only used by germinating plant embryos but also eaten by animals, including humans. Development of the embryo and endosperm following fertilization is possible because of the constant flow of nutrients into the developing seed from the parent plant. Cell divisions of the fertilized egg to form a multicellular embryo proceed in several ways in flowering plants. We describe eudicot embryonic development; monocot embryonic development is similar in the early stages.

The two cells (basal cell and apical cell) that are formed as a result of the first division of the fertilized egg establish polarity, or direction, in the embryo. The large basal cell (located toward the outside of the ovule) typically develops into a suspensor, an embryonic tissue that anchors the developing embryo and aids in nutrient uptake from the endosperm. The apical cell (toward the inside of the ovule) develops into the plant embryo. Scientists are currently studying molecular differences between the apical cell and basal cell to determine the initial molecular signals that are involved in establishing polarity. Initially, the apical cell divides to form a small cluster of cells, called a proembryo (FIG. 37-8). As cell division continues, a sphere of cells, often called a globular embryo, develops. Cells begin to develop into specialized tissues during this stage. When the eudicot embryo starts to develop its two cotyledons (seed leaves), it has two lobes and resembles a heart; this is often called the heart stage. During the torpedo stage, the embryo continues to grow as the cotyledons elongate. As the embryo enlarges, it often curves back on itself and crushes the suspensor.

Anther

The mature seed contains an embryonic plant and storage materials A mature seed contains an embryonic plant and food (stored in the cotyledons or endosperm), surrounded by a tough, protective seed coat derived from the integuments, which are the outermost layers of an ovule. The seed, in turn, is enclosed within a fruit.

Pollen grains

4

Stigma 1

Generative cell

Pollination occurs.

Double fertilization occurs.

Pollen tube

Tube cell Style

2

Pollen grain

Pollen tube grows through style to ovule in ovary. 3 nuclei (sperm cell and two polar nuclei) fuse to form endosperm.

Ovary Ovule (containing female gametophyte) Integuments

2 polar nuclei Meiosis and mitosis

Megasporocyte Female gametophyte

Sperm cells

Egg 3

Tube nucleus

FIGURE 37-7 Animated Pollination, pollen tube growth, and double fertilization

Two sperm cells move down pollen tube and enter ovule.

Sperm cell and egg fuse to form zygote.

Endosperm

Proembryo developing within ovule

Torpedo stage of embryo

Emerging cotyledons

Globular embryo

Heart-shaped embryo

Suspensor

Stem tip

Root tip

Suspensor Basal cell

1

Proembryo. This is the earliest multicellular stage of the embryo.

2

Globular stage embryo. As cell division continues, the embryo becomes a ball of cells.

3

Heart stage embryo. As two cotyledons begin to emerge, the embryo is shaped like a heart.

4

Torpedo stage embryo. Cotyledons continue to elongate, forming the torpedo stage.

Developing seed coat Maturing embryo

Stem apical meristem

Mature seeds

Two cotyledons Root apical meristem Remains of endosperm

5

Heart-shaped fruit containing many seeds

Maturing embryo within seed. Food originally stored in endosperm has been almost completely depleted during embryonic growth and development, and most of the food for the embryonic plant is now stored in the cotyledons.

6

FIGURE 37-8 Embryonic development in shepherd’s purse (Capsella bursa-pastoris)

Cotyledon

Radicle

James Mauseth, University of Texas

The ovule is shown apart from the ovary in parts ● 1 through ● 5.

The mature embryo within the seed consists of a short embryonic root, or radicle; an embryonic shoot; and one or two seed leaves, or cotyledons (FIG. 37-9). Monocots have a single cotyledon, whereas eudicots have two. The short portion of the embryonic shoot connecting the radicle to one or two cotyledons is the hypocotyl. The shoot apex, or terminal bud, located above the point of attachment of the cotyledon(s) is the plumule. After the radicle, hypocotyl, cotyledon(s), and plumule have formed, the young plant’s development is arrested, usually by desiccation (drying out) or dormancy (a temporary state of arrested physiological activity). When conditions are right for continuing the developmental program, the seed germinates, and the embryo resumes growth. Many seeds are known to remain dormant for decade or so, but only a few instances of germination after hundreds of years of dormancy have been scientifically documented. In 2008 a team

Fruit. Longitudinal section through a fruit of shepherd’s purse reveals numerous tiny seeds, each containing a mature embryo. Each seed developed from an ovule.

Foliage leaf

Plumule Hypocotyl

FIGURE 37-9 Seed structure A bean seed has been dissected—its seed coat and one of its two cotyledons were removed—to show the radicle, hypocotyl, remaining cotyledon, and plumule with its foliage leaf at the shoot apex. This particular seed had begun germinating, so its radicle is larger than in an ungerminated seed.

Seed

Berry (simple fruit) A simple, fleshy fruit in which the fruit wall is soft throughout. Tomato (Lycopersicon lycopersicum)

Fused fruit wall and seed coat

Caryopsis (simple fruit) A simple, dry fruit in which the fruit wall is fused to the seed coat. Wheat (Triticum) Single seed

Single seed Drupe (simple fruit) A simple, fleshy fruit in which the inner wall of the fruit is a hard stone. Peach (Prunus persica)

Seed coat

Woody fruit wall Follicle (simple fruit) A simple, dry fruit that splits open along one suture to release its seeds; fruit is formed from ovary that consists of a single carpel.

Seed

Split-open suture

Seed

Capsule (simple fruit) A simple, dry fruit that splits open along two or more sutures or pores to release its seeds; fruit is formed from ovary that consists of two or more carpels. Iris (Iris)

Nut (simple fruit) A simple, dry fruit that has a stony wall, is usually large, and does not split open at maturity. Oak (Quercus)

Single seed Aggregate fruit A fruit that develops from a single flower with several to many pistils (i.e., carpels are not fused into a single pistil). Blackberry (Rubus)

Seed

Milkweed (Asclepias syriaca)

Legume (simple fruit) A simple, dry fruit that splits open along two sutures to release its seeds; fruit is formed from ovary that consists of a single carpel. Green bean (Phaseolus vulgaris)

Achene (simple fruit) A simple, dry fruit in which the fruit wall is separate from the seed coat. Sunflower (Helianthus annuus)

Scale-covered cup

Single seed inside stone

Seed

Fruit wall

Multiple fruit A fruit that develops from the ovaries of a group of flowers. Mulberry (Morus)

Seed

Enlarged floral tube Ovary wall

FIGURE 37-10 Fruit types Fruits are botanically classified into four groups—simple, aggregate, multiple, and accessory— based on their structure and mechanism of seed dispersal.

Seed

Accessory fruit A fruit composed primarily of nonovarian tissue (such as the receptacle or floral tube). Apple (Malus sylvestris)

of biologists from Israel and Switzerland successfully germinated a date seed (Phoenix dactylifera) that was radiocarbon dated at about 2000 years old. Before this report, the oldest documented instance of germination was a 1200-year-old lotus seed (Nelumbo nucifera). Because the embryonic plant is nonphotosynthetic, it must be nourished during germination until it becomes photosynthetic and therefore self-sufficient. The cotyledons of many plants function as storage organs and become large, thick, and fleshy as they absorb the food reserves (starches, oils, and proteins) initially produced as endosperm. Seeds that store nutrients in cotyledons have little or no endosperm at maturity. Examples of such seeds are peas, beans, squashes, sunflowers, and peanuts. Other plants—wheat and corn, for example—have thin cotyledons that function primarily to help the young plant digest and absorb food stored in the endosperm.

Fruits are mature, ripened ovaries After double fertilization takes place within the ovule, the ovule develops into a seed, and the ovary surrounding it develops into a fruit. For example, a pea pod is a fruit, and the peas within it are seeds. A fruit may contain one or more seeds; some orchid fruits contain several thousand to a few million seeds! Fruits provide protection for the enclosed seeds and sometimes aid in their dispersal. There are several types of fruits; their differences result from variations in the structure or arrangement of the flowers from which they were formed. The four basic types of fruits—simple fruits, aggregate fruits, multiple fruits, and accessory fruits—are summarized in FIGURE 37-10. Most fruits are simple fruits. A simple fruit develops from a single ovary, which may consist of a single carpel or several fused carpels. At maturity, simple fruits may be fleshy or dry. Two examples of simple, fleshy fruits are berries and drupes. A berry is a fleshy fruit that has soft tissues throughout and contains few to many seeds; a blueberry is a berry, as are grapes, cranberries, bananas, and tomatoes. Many so-called “berries” do not fit the botanical definition. Strawberries, raspberries, and mulberries, for example, are not berries; these three fruits are discussed shortly.

A drupe is a simple, fleshy or fibrous fruit that contains a hard stone (pit) surrounding a single seed. Examples of drupes include peaches, plums, olives, and almonds. The almond shell is actually the stone, which remains after the rest of the fruit has been removed. Many simple fruits are dry at maturity; some of these split open, usually along seams, called sutures, to release their seeds. A milkweed pod is an example of a follicle, a simple, dry fruit that develops from a single carpel and splits open along one suture to release its seeds. A legume is a simple, dry fruit that develops from a single carpel and splits open along two sutures. Pea pods are legumes, as are green beans, although both are generally harvested before the fruit has dried out and split open. Pea seeds are usually removed from the fruit and consumed, whereas in green beans the entire fruit and seeds are eaten. A capsule is a simple, dry fruit that develops from two or more fused carpels and splits open along two or more sutures or pores. Iris, poppy, and cotton fruits are capsules. Other simple, dry fruits, such as caryopses (sing., caryopsis), or grains, do not split open at maturity. Each caryopsis contains a single seed. Because the seed coat is fused to the fruit wall, a caryopsis looks like a seed rather than a fruit. Kernels of corn and wheat are fruits of this type. An achene is similar to a caryopsis in that it is simple and dry, does not split open at maturity, and contains a single seed. However, the seed coat of an achene is not fused to the fruit wall. Instead, the single seed is attached to the fruit wall at one point only, permitting an achene to be separated from its seed. The sunflower fruit is an example of an achene. One can peel off the fruit wall (the shell) to reveal the sunflower seed within. Nuts are simple, dry fruits that have a stony wall and do not split open at maturity. Unlike achenes, nuts are usually large, single seeded, and often derived from a compound ovary. Examples of nuts include chestnuts, acorns, and hazelnuts. Many so-called “nuts” do not fit the botanical definition. Peanuts and Brazil nuts, for example, are seeds, not nuts. Aggregate fruits are a second main type of fruit. An aggregate fruit is formed from a single flower that contains several to many separate (free) carpels (FIG. 37-11). After fertilization, each ovary from each individual carpel enlarges. As they enlarge, the ovaries

Stigmas and styles Stamens Remnants of stigmas and styles Ovaries (in a) become tiny drupes (in b) Receptacle Remnants of stamens

Cengage

Sepal

Petal Ovaries of separate carpels

(a)

(b)

(c)

FIGURE 37-11 An aggregate fruit (a) Cutaway view of a blackberry flower (Rubus), showing the many separate carpels in the center of the flower. (b) A developing blackberry fruit is an aggregate of tiny drupes. The little “hairs” on the blackberry are remnants of stigmas and styles. (c) Developing blackberry fruits at various stages of maturity.

may fuse to form a single fruit. Raspberries, blackberries, and magnolia fruits are examples of aggregate fruits. A third type is the multiple fruit, formed from the ovaries of many flowers that grow in proximity on a common floral stalk. The ovary from each flower fuses with nearby ovaries as it develops and enlarges after fertilization. Pineapples, figs, and mulberries are multiple fruits. Accessory fruits are the fourth type. They differ from other fruits in that plant tissues in addition to ovary tissue make up the fruit. For example, the edible portion of a strawberry is the red, fleshy receptacle. Apples and pears are also accessory fruits; the outer part of each of these fruits is an enlarged floral tube, consisting of receptacle tissue along with portions of the calyx, that surrounds the ovary (FIG. 37-12).

Seed dispersal is highly varied Wind, animals, water, and explosive dehiscence disperse the various seeds and fruits of flowering plants (FIG. 37-13). Effective methods of seed dispersal have made it possible for certain plants to expand their geographic range. In some cases the seed is the actual agent of dispersal, whereas in others the fruit performs this role. In tumbleweeds, such as Russian thistle, the entire plant is the agent of dispersal because it detaches and blows across the ground, scattering seeds as it bumps along. Tumbleweeds are lightweight and are sometimes blown many kilometers by the wind. Wind is responsible for seed dispersal in many plants. Plants such as maple trees have winged fruits adapted for wind dispersal. Light, feathery plumes enable other seeds or fruits to be transported by wind, often for considerable distances. Both dandelion fruits and milkweed seeds have this type of adaptation.

Some plants have special structures that aid in dispersal of their seeds and fruits by animals. The spines and barbs of burdock burs and similar fruits catch in animal fur and fall off as the animal moves about. Fleshy, edible fruits are also adapted for animal dispersal. Birds, bats, primates, grazing ruminants, and ants are common dispersal agents. These animals are attracted to the fruit by its color, location, odor, and taste. As the animal eats these fruits, it either discards or swallows the seeds. Many seeds that are swallowed have thick seed coats and are not digested; instead, they pass through the digestive tract and are deposited in the animal’s feces some distance from the parent plant. In fact, some seeds do not germinate unless they have passed through an animal’s digestive tract; the animal’s digestive juices probably aid germination by helping to break down the seed coat. Some edible fruits apparently contain chemicals that function as laxatives to speed seeds through an animal’s digestive tract; the less time these seeds spend in the gut, the more likely they will germinate. Animals such as squirrels and many bird species also help disperse acorns and other fruits and seeds by burying them for winter use. Many buried seeds are never used by the animal and germinate the following spring. Ants collect the seeds of many plants and take them underground to their nests. Ants disperse and bury seeds for hundreds of plant species in almost every terrestrial environment, from northern coniferous forests to tropical rain forests to deserts. Both ants and flowering plants benefit from their association. The ants ensure the reproductive success of the plants whose seeds they bury, and the plants supply food to the ants. A seed that is collected and taken underground by ants often contains a special structure called an elaiosome, or oil body, that protrudes from the

Petal Stigma

Sepals

Stamens Style

Ovary Floral tube Ovule (in a) becomes seed (in b)

Sepal

(a)

FIGURE 37-12 An accessory fruit (a) Note the floral tube surrounding the ovary in the pear flower (Pyrus communis). This tube becomes the major edible portion of the pear. (b) Longitudinal section through a pear showing the fruit tissue, which derives from both the floral tube and the ovary.

(b)

DPA / Dembinsky Photo Associates

© Gary Meszaros/ Visuals Unlimited

John Serrao / Visuals Unlimited

(a) The feathery plumes of milkweed (Asclepias syriaca ) seeds make them buoyant for dispersal by wind.

(c) Fleshy fruits such as blackberries (Rubus) are eaten by animals such as this white-footed mouse. The seeds are frequently swallowed whole and pass unharmed through the animal’s digestive tract.

Marion Lobstein

(b) Burdock (Arctium minus) burs (the hooked fruits) are carried away from the parent plant after sticking to bird feathers, mammal fur, or human clothing.

(d) Dispersal of seeds by ants. The brown part of each bloodroot (Sanguinaria canadensis) seed is the seed proper, and the white part is the elaiosome, or oil body. The seeds have been placed along the midvein of an oak leaf to indicate scale.

FIGURE 37-13 Seed (and fruit) dispersal

(e) (Left) An intact fruit of bitter cress (Cardamine pratensis), before it has opened. (Right) The fruit splits open with explosive force, flinging the seeds some distance from the plant.

seed. Elaiosomes are a nutritious food for ants, which carry seeds underground before removing the elaiosome. Once an elaiosome is removed from a seed, the ants discard the undamaged seed in an underground refuse pile, which happens to be rich in organic material (such as ant droppings and dead ants) and minerals (inorganic nutrients) required by young seedlings. Thus, ants not only bury the seeds away from animals that might eat them but also place the seeds in rich soil that is ideal for germination and seedling growth. The coconut is an example of a fruit adapted for dispersal by water. The coconut has air spaces that make it buoyant and capable of being carried by ocean currents for thousands of kilometers. When it washes ashore, the seed may germinate and grow into a coconut palm tree. Some seeds are not dispersed by wind, animals, or water. Such seeds are found in fruits that use explosive dehiscence, in which the fruit bursts open suddenly and quite often violently, forcibly discharging its seeds. These fruits burst open due to pressure caused by differences in turgor (hydrostatic pressure) as the cells in the fruit dry out. The fruits of plants such as touch-me-not and bitter cress split open so explosively that seeds are scattered a meter or more.

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What is the difference between pollination and fertilization? Which process occurs first? What are the main stages of eudicot embryonic development? What is a fruit? How are the following fruits distinguished: simple, aggregate, multiple, and accessory? What are some characteristics of animal-dispersed seeds and fruits?

37.4 GERMINATION AND EARLY GROWTH ■ ■ LEARNING OBJECTIVE

and others degraded. Therefore, water is an absolute requirement for germination. The absorption of water by a dry seed is imbibition. As a seed imbibes water, it often swells to several times its original, dry size (FIG. 37-14). Cells imbibe water by adhesion of water onto and into materials such as cellulose, pectin, and starches within the seed. Germination and subsequent growth require a great deal of energy. Because young plants obtain this energy by converting the energy of food molecules stored in the seed’s endosperm or cotyledons to ATP during aerobic respiration, much oxygen is usually needed during germination. (Some plants, such as rice, grow in flooded soil where oxygen is absent and carry out alcohol fermentation during the early stages of germination and seedling growth.) Temperature is another environmental factor that affects germination. Each species has an optimal, or ideal, temperature at which the germination percentage is highest. For most plants, the optimal germination temperature is between 25°C and 30°C (77°F and 86°F). Some seeds, such as those of apples, require prolonged exposure to low temperatures before their seeds break dormancy and germinate. Some of the environmental factors needed for germination help ensure the survival of the young plant. The requirement of a prolonged low-temperature period ensures that seeds adapted to temperate climates germinate in the spring rather than the fall. Some plants, especially those with tiny seeds, such as lettuce, require light for germination. A light requirement ensures that a tiny seed germinates only if it is close to the soil surface. If such a seed were to germinate several centimeters below the soil surface, it might not have enough food reserves to grow to the surface. But if this light-dependent seed remains dormant until the soil is disturbed and it is brought to the surface, it has a much greater likelihood of survival.

Some seeds do not germinate immediately A mature seed is often dormant and may not germinate immediately even if growing conditions are ideal. In certain seeds, internal factors, which are under genetic control, prevent germination even

9 Summarize the influence of internal and environmental factors on the

Pollination and fertilization are followed by seed and fruit development in flowering plants. Each seed develops from an ovule and contains an embryonic plant and food to provide nourishment for the embryo during germination. Germination—the process of a seed sprouting—and the growth of young seedlings into mature plants are aspects of growth and development. Within a given species, the precise requirements for seed germination represent evolutionary adaptations that protect the young seedlings from adverse environmental conditions. Environmental cues, such as the presence of water and oxygen, proper temperature, and sometimes the presence of light penetrating the soil surface, influence whether or not a seed germinates. No seed germinates unless it has absorbed water. The embryo in a mature seed is dehydrated, and a watery environment is necessary for active metabolism. When a seed germinates, its metabolic machinery is turned on, and numerous materials are synthesized

Marion Lobstein

germination of seeds.

FIGURE 37-14 Imbibition Pinto bean seeds (Phaseolus vulgaris) before imbibition (left) and after (right). Dry seeds imbibe water before they germinate.

when all external conditions are favorable. Many seeds are dormant because certain chemicals are present or absent or because the seed coat restricts germination. For example, the seeds of many desert plants contain high concentrations of abscisic acid (discussed in Chapter 38), which inhibits germination under unfavorable conditions. Abscisic acid is washed out only when rainfall is sufficient to support the plant’s growth after the seed germinates. Some seeds, such as certain legumes, have extremely hard, thick seed coats that prevent water and oxygen from entering, thereby inducing dormancy. Scarification, the process of scratching or scarring the seed coat (physically with a knife or chemically with an acid) before sowing it, induces germination in these plants. Scarification in nature occurs, for example, when these seeds pass through the digestive tracts of animals or when the seed coats are partially digested by soil bacteria.

Eudicots and monocots exhibit characteristic patterns of early growth Once conditions are right for germination, the first part of the plant to emerge from the seed is the radicle, or embryonic root. As the root grows and forces its way through the soil, it encounters considerable friction from soil particles. A root cap protects the delicate apical meristem of the root tip. The shoot (the stem with its leaves and reproductive structures) is next to emerge from the seed. Stem tips are not protected by a structure comparable to a root cap, but plants have ways to protect the delicate stem tip as it grows through the soil to the surface. The stem of a bean seedling (a eudicot), for instance, curves over to form a hook so that the stem tip and cotyledons are actually pulled up through the soil (FIG. 37-15a). Corn and other grasses (monocots) have a special sheath of cells called a coleoptile that surrounds and protects the young shoot (FIG. 37-15b). First, the coleoptile pushes up through the soil, and then the leaves and stem grow through the tip of the coleoptile.

Review ■ ■

What factors influence the germination of seeds? Why are plant growth and development so sensitive to environmental cues?

37.5 ASEXUAL REPRODUCTION IN FLOWERING PLANTS ■ ■ LEARNING OBJECTIVES 10 Explain how the following structures may be used to propagate plants asexually: rhizomes, tubers, bulbs, corms, stolons, plantlets, and suckers.

11 Define apomixis.

Flowering plants have many kinds of asexual reproduction, several of which involve modified stems: rhizomes, tubers, bulbs, corms, and stolons. A rhizome is a horizontal underground stem that may or may not be fleshy. Fleshiness indicates that the rhizome is used for storing food materials, such as starch (FIG. 37-16a). Although rhizomes resemble roots, they are really stems, as indicated by the

First foliage leaf

Cotyledons (two) Hook Coleoptile

Seed coat

Branch root Primary root

Adventitious root Endosperm Cotyledon (one) Branch root Primary root

(a) Common bean, a eudicot. The hook in the stem protects the delicate stem tip as it grows through the soil. Once the shoot emerges from the soil, the hook straightens.

(b) Corn, a monocot. The coleoptile, a sheath of cells, emerges first from the soil. The shoot and leaves grow through the middle of the coleoptile.

FIGURE 37-15 Animated Germination and seedling growth

presence of scalelike leaves, buds, nodes, and internodes. (Roots have none of these features.) Rhizomes frequently branch in different directions. Over time, the old portion of the rhizome dies, and the two branches eventually separate to become distinct plants. Irises, bamboos, ginger, and many grasses are examples of plants that reproduce asexually by forming rhizomes. Some rhizomes produce greatly thickened ends called tubers, which are fleshy underground stems enlarged for food storage. When the attachment between a tuber and its parent plant breaks, often as a result of the death of the parent plant, the tuber grows into a separate plant. Potatoes and elephant’s ear (Caladium) are examples of plants that produce tubers (FIG. 37-16b). The “eyes” of a potato are axillary buds, evidence that the tuber is an underground stem rather than a storage root such as a sweet potato or carrot. A bulb is a modified underground bud in which fleshy storage leaves are attached to a short stem (FIG. 37-16c). A bulb is globose (round) and covered by paperlike bulb scales, which are modified leaves. It frequently forms axillary buds that develop into small daughter bulbs (bulblets). These new bulbs are initially attached to the parent bulb, but when the parent bulb dies and rots away, each daughter bulb can become established as a separate plant. Lilies, tulips, onions, and daffodils are some plants that form bulbs. A corm is a short, erect underground stem that superficially resembles a bulb (FIG. 37-16d). Unlike the bulb, whose food is stored in underground leaves, the corm’s storage organ is a thickened underground stem covered by papery scales (modified leaves). Axillary buds frequently give rise to new corms; the death of the parent corm separates these daughter corms, which then become established as separate plants. Familiar garden plants that produce corms include crocus, gladiolus, and cyclamen.

Rhizome

Rhizome Adventitious roots

Bulb

Stem

Tuber

Adventitious roots

Roots

(a) Rhizome. Irises have horizontal underground stems called rhizomes. New aerial shoots arise from buds that develop on the rhizome.

Fleshy leaves

(b) Tuber. Potato plants form rhizomes, which enlarge into tubers (the potatoes) at the ends.

(c) Bulb. A bulb is a short underground stem to which overlapping, fleshy leaves are attached; most of the bulb consists of leaves.

Axillary bud

Leaf scars Corm (modified stem) Old corm (last year’s)

New shoot

Adventitious roots

(d) Corm. A corm is an underground stem that is almost entirely stem tissue surrounded by a few papery scales.

Scale leaf (at node)

Adventitious roots

Stolon (runner)

(e) Stolon. Strawberries reproduce asexually by forming stolons, or runners. New plants (shoots and roots) are produced at every other node.

FIGURE 37-16 Modified stems

Stolons, or runners, are horizontal, aboveground stems that grow along the surface and have long internodes (FIG. 37-16e). Buds develop along the stolon, and each bud gives rise to a new shoot that roots in the ground. When the stolon dies, the daughter plants live separately. The strawberry plant produces stolons. Some plants form detachable plantlets (small plants) in notches along their leaf margins. Kalanchoe, whose common name is “mother of thousands,” has meristematic tissue that gives rise to an individual plantlet at each notch in the leaf (FIG. 37-17). When these plantlets reach a certain size, they drop to the ground, root, and grow.

Some plants reproduce asexually by producing suckers, aboveground shoots that develop from adventitious buds on roots (FIG. 37-18). Each sucker grows additional roots and becomes an independent plant when the parent plant dies. Examples of plants that form suckers include black locust, pear, apple, cherry, blackberry, and aspen. A quaking aspen (Populus tremuloides) colony in the Wasatch Mountains of Utah contains at least 47,000 tree trunks formed from suckers that can be traced back to a single individual; this massive “organism” occupies almost 43 hectares (106 acres). Some weeds, such as field bindweed, produce many suckers. These plants are difficult to control because pulling the plant out of the

Jerome Wexler/Photo Researchers, Inc.

FIGURE 37-17 Plantlets The “mother of thousands” (Kalanchoe) produces detachable plantlets along the margins of its leaves. The young plantlets drop off and root in the ground.

of asexual reproduction in which the offspring are maternal clones. For example, an embryo may develop from a diploid cell in the ovule rather than from a diploid zygote that forms from the union of two haploid gametes. Because there is no fusion of gametes, the embryo is virtually genetically identical to the maternal genotype. However, the advantage of apomixis over other methods of asexual reproduction is that the seeds and fruits produced by apomixis can be dispersed by methods associated with sexual reproduction. Apomixis, which occurs in various species of more than 40 angiosperm families, is often found in plants that are polyploid with varying degrees of sterility. Apomixis allows these sterile plants to reproduce and therefore survive. Examples of plants that can reproduce by apomixis include dandelions, citrus trees, mango, blackberries, garlic, and certain grasses. Plant biologists are actively engaged in identifying the genes involved in asexual reproduction by apomixis. In 2008, scientists in India reported in the journal Nature that they had identified a mutation of a single gene in Arabidopsis that results in diploid female gametes, a first step toward apomixis. If researchers can transfer this or similar genes to crop plants that possess superior traits such as high yield, they may be able to develop apomictic crop plants whose traits do not get lost or diluted during the genetic shuffling of sexual reproduction.

Review ■ ■

Original plant

■

How are rhizomes and tubers involved in asexual reproduction? How are corms and bulbs similar? How are they different? What is apomixis? How is the ability to reproduce by apomixis advantageous?

Suckers

37.6 A COMPARISON OF SEXUAL AND ASEXUAL REPRODUCTION

Sucker

■ ■ LEARNING OBJECTIVE 12 State the differences between sexual reproduction and asexual reproRoot system

FIGURE 37-18 Suckers in various stages of development All the suckers are connected to one another through the root system.

soil seldom removes all the roots, which can grow as deep as 3 m (10 ft). In fact, in response to wounding, the roots produce additional suckers, which can be a considerable nuisance to humans.

Apomixis is the production of seeds without the sexual process Some flowering plants produce embryos in seeds without meiosis and the fusion of gametes. Seed production by apomixis is a form

duction, and discuss the evolutionary advantages and disadvantages of each.

Sexual reproduction and asexual reproduction are suited for different environmental circumstances. As you know, sexual reproduction results in offspring that are genetically different from the parents; that is, the parental genotypes are not preserved in the offspring. This genetic diversity of offspring may be selectively advantageous, particularly in an unstable, or changing, environment. (We define environment broadly to include all external conditions, both living and nonliving, that affect an organism.) If a plant species that reproduces sexually (and is therefore genetically diverse) is exposed to increasing annual temperatures as a result of global warming, for example, some of the individuals may be more fit than the parents or other offspring. The genetic diversity from sexual reproduction may also let the individuals of a species exploit new environments, thereby expanding their range.

You have seen that asexual reproduction results in offspring that are virtually genetically identical to the parent; that is, the parental genotype is preserved. Assuming the parent is well adapted to its environment (that is, the parent has a favorable combination of alleles), this genetic similarity may be selectively advantageous if the environment remains stable (unchanging) for several generations. None of the offspring of asexual reproduction is more fit than the parent, but neither is any less fit. Despite the apparent advantages of asexual reproduction, most plant species whose reproduction is primarily asexual occasionally reproduce sexually. Even in what appears to be a stable environment, plants are exposed to changing selective pressures, such as changes in the number and kinds of herbivores and parasites, the availability of food, competition from other species, and climate. Sexual reproduction permits species whose reproduction is primarily asexual to increase their genetic variability so at least some individuals are adapted to the changing selective pressures in a stable environment.

Sexual reproduction has some disadvantages

tion is a costly form of reproduction. Sexual reproduction requires both male and female gametes, which must meet for reproduction to occur. The many adaptations of flowers for different modes of pollination represent one cost of sexual reproduction. Sexual reproduction produces some individuals with genotypes that are well-adapted to the environment, but it also produces some individuals that are less well-adapted. Therefore, sexual reproduction is usually accompanied by high death rates among offspring, particularly when selective pressures are strong. (As discussed in Chapter 18, however, this aspect of sexual reproduction is an important part of evolution by natural selection.) Every biological process involves trade-offs, and sexual reproduction is no exception. Although sexual reproduction has its costs, it also has adaptive advantages over asexual reproduction.

Review How does sexual reproduction in plants differ from asexual reproduction? What is an adaptive advantage of sexual reproduction? Why is sexual reproduction considered costly?

■

■

Although the genetic diversity produced by sexual reproduction is advantageous to a species’ long-term survival, sexual reproduc-

■ ■

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

37.1 (page 782)

■

1 Describe the functions of each part of a flower. ■

■

■

Sepals cover and protect the flower parts when the flower is a bud. Petals play an important role in attracting animal pollinators to the flower. Stamens produce pollen grains. Each stamen consists of a thin stalk (the filament) attached to a saclike structure (the anther). The carpel is the female reproductive unit. A pistil may consist of a single carpel or a group of fused carpels. Each pistil has three sections: a stigma, on which the pollen grains land; a style, through which the pollen tube grows; and an ovary that contains one or more ovules.

An egg and two polar nuclei, along with several other nuclei, are formed in the ovule. Both egg and polar nuclei participate directly in fertilization. Watch plant reproduction unfold by clicking on the figure in CengageNOW.

37.2 (page 784) 3 Compare the evolutionary adaptations that characterize flowers pollinated in different ways (by insects, birds, and bats). ■ Flowers pollinated by insects are often yellow or blue and have a scent. Bird-pollinated flowers are often yellow, orange, or red and do not have a strong scent. Bat-pollinated flowers often have white petals and are scented. 4 Define coevolution and give examples of ways that plants and their animal pollinators have affected one another’s evolution. Coevolution is reciprocal adaptation caused by two different spe■ cies (such as flowering plants and their animal pollinators) forming an interdependent relationship and affecting the course of each other’s evolution. For example, flowers with large, showy petals and scent have evolved in some plants, whereas hairy bodies that catch and hold sticky pollen grains have evolved in bees.

37.3 (page 789) 5 Distinguish between pollination and fertilization. ■

Learn more about flower structure by clicking on the figure in CengageNOW.

2 Identify where eggs and pollen grains are formed within the flower. ■

Pollen grains form within pollen sacs in the anther.

■

Pollination is the transfer of pollen grains from anther to stigma. After pollination, fertilization, the fusion of gametes, occurs. Flowering plants undergo double fertilization. The egg fuses with one sperm cell, forming a zygote (fertilized egg) that eventually develops into a multicellular embryo in the seed. The two polar nuclei fuse with the second sperm cell, forming a triploid nutritive tissue called endosperm.

Learn more about double fertilization by clicking on the figures in CengageNOW.

6 Trace the stages of embryo development in flowering plants, and list and define the main parts of seeds. ■ A eudicot embryo develops in the seed in an orderly fashion, from proembryo to globular embryo to the heart stage to the torpedo stage. ■ A mature seed contains both a young plant embryo and nutritive tissue—stored in the endosperm or in the cotyledons (seed leaves)—for use during germination. A tough, protective seed coat surrounds the seed. Watch embryonic development in flowering plants by clicking on the figure in CengageNOW.

7 Explain the relationships among the following: ovules, ovaries, seeds, and fruits. Ovules are structures with the potential to develop into seeds, whereas ovaries are structures with the potential to develop into fruits. Seeds are enclosed within fruits, which are mature, ripened ovaries. 8 Distinguish among simple, aggregate, multiple, and accessory fruits. ■ Simple fruits develop from a single ovary that consists of one carpel or several fused carpels. Aggregate fruits develop from a single flower with many separate ovaries. Multiple fruits develop from the ovaries of many flowers growing in close proximity on a common axis. In accessory fruits, the major part of the fruit consists of tissue other than ovary tissue. ■

37.4 (page 796) 9 Summarize the influence of internal and environmental factors on the germination of seeds. Germination is the process of seed sprouting. Internal factors affecting whether a seed germinates include the maturity of the embryo; the presence or absence of chemical inhibitors; and the presence or absence of hard, thick seed coats. ■ External environmental factors that may affect germination include requirements for oxygen, water, temperature, and light. ■

Learn more about germination and seedling growth by clicking on the figure in CengageNOW.

37.5 (page 797) 10 Explain how the following structures may be used to propagate plants asexually: rhizomes, tubers, bulbs, corms, stolons, plantlets, and suckers. A rhizome is a horizontal underground stem. A tuber is a fleshy ■ underground stem enlarged for food storage. A bulb is a modified underground bud with fleshy storage leaves attached to a short stem. A corm is a short, erect underground stem covered by papery scales. A stolon is a horizontal aboveground stem with long internodes. ■ Some leaves have meristematic tissue along their margins and give rise to detachable plantlets. ■ Roots may develop adventitious buds that develop into suckers. Suckers produce additional roots and may give rise to new plants. 11 Define apomixis. ■ Apomixis is the production of seeds and fruits without sexual reproduction.

37.6 (page 799) 12 State the differences between sexual reproduction and asexual reproduction, and discuss the evolutionary advantages and disadvantages of each. ■ Sexual reproduction involves the union of two gametes; the offspring produced by sexual reproduction are genetically variable. Asexual reproduction involves the formation of offspring without the fusion of gametes; the offspring are virtually genetically identical to the single parent. ■ Genetic diversity among offspring produced by sexual reproduction may let certain individuals survive in a changing environment or exploit new environments. Sexual reproduction is costly because both male and female gametes must be produced and must meet. ■ Genetic similarity among offspring produced by asexual reproduction may be advantageous if the environment is stable. Most plant species whose reproduction is primarily asexual occasionally reproduce sexually, thereby increasing their genetic variability.

T E S T YO U R U N D E R S TA N D I N G 1. The normal order of whorls from the flower’s periphery to the center is (a) sepals, petals, carpels, stamens (b) stamens, carpels, sepals, petals (c) sepals, petals, stamens, carpels (d) petals, carpels, stamens, sepals (e) carpels, stamens, petals, sepals

5. The process of _____ in flowering plants involves one sperm cell fusing with an egg cell and one sperm cell fusing with two polar nuclei. (a) coevolution (b) germination (c) double fertilization (d) apomixis (e) pollination

2. A pistil consists of (a) stigma, style, and stamen (b) anther and filament (c) sepal and petal (d) stigma, style, and ovary (e) radicle, hypocotyl, and plumule

6. The nutritive tissue in the seeds of flowering plants that is formed from the union of a sperm cell and two polar nuclei is called the (a) plumule (b) endosperm (c) cotyledon (d) hypocotyl (e) radicle

3. The transfer of pollen grains from anther to stigma is (a) fertilization (b) double fertilization (c) pollination (d) germination (e) apomixis 4. The observation that insects with long mouthparts pollinate long, tubular flowers and insects with short mouthparts pollinate flowers with short corollas is explained by (a) coevolution (b) germination (c) double fertilization (d) apomixis (e) explosive dehiscence

7. After fertilization, the ovule develops into a _____ and the ovary develops into a _____. (a) fruit; seed (b) seed; fruit (c) calyx; corolla (d) corolla; calyx (e) follicle; legume 8. In plants that lack endosperm in their mature seeds, the cotyledons function to (a) enclose and protect the seed (b) aid in seed dispersal (c) serve as an absorptive embryonic root (d) store food reserves (e) attach the embryo within the ovule

9. _____ fruits develop from many ovaries of a single flower, whereas _____ fruits develop from the ovaries of many separate flowers. (a) multiple; accessory (b) simple; accessory (c) aggregate; multiple (d) accessory; aggregate (e) simple; multiple 10. A horizontal underground stem that may or may not be fleshy and that is often specialized for asexual reproduction is called a (a) stolon (b) bulb (c) corm (d) rhizome (e) tuber

11. Place the following events in the correct order. 1. pollen tube grows into ovule 2. insect lands on flower to drink nectar 3. embryo develops within the seed 4. fertilization occurs 5. pollen carried by insect contacts stigma (a) 2, 5, 1, 4, 3 (b) 1, 4, 2, 5, 3 (c) 3, 2, 5, 1, 4 (d) 5, 1, 3, 4, 2 (e) 2, 5, 4, 3, 1 12. Label the following diagram and give the function of each part. (Refer to Figure 37-1b to check your answers.)

CRITICAL THINKING 1. Sketch the kinds of flowers that form simple, aggregate, multiple, and accessory fruits. 2. Using what you have learned in this chapter, speculate whether it is more likely that offspring of asexual reproduction develop in close proximity to or widely dispersed from the parent plant. Explain your reasoning. How could you design an experiment to test your hypothesis? 3. Which type of reproduction, sexual or asexual, might be more beneficial in the following circumstances, and why: (a) a perennial (plant that lives more than two years) in a stable environment; (b) an annual (plant that lives one year) in a rapidly changing environment; and (c) a plant adapted to an extremely narrow climate range?

4. Using what you have learned in this chapter, offer an explanation of why telephone poles and wires strung across grassy fields or plains often have tree seedlings growing under them. 5. EVOLUTION LINK. Is seed dispersal by ants an example of coevolution? Why or why not? 6. SCIENCE, TECHNOLOGY, AND SOCIETY. What is the value to society of research on genes that control flowering? Additional questions are available in CengageNOW at www.cengage.com/ login.

38

Plant Developmental Responses to External and Internal Signals

Dwight R. Kuhn

Black-eyed Susan. This plant (Rudbeckia hirta) produces flowers in response to the shortening nights of spring and early summer.

KEY CONCEPTS

T

38.1 A plant may respond to an external stimulus, such as

is genetic. If the genes required for development of a particular trait, such

light, gravity, or touch, by a directional growth response, or tropism. Study of tropisms has helped biologists elucidate links between the external environment and internal signals such as hormones.

as the shape of a leaf, the color of a flower, or the type of root system,

38.2 Hormones are chemical signals responsible for coordinating and regulating many aspects of plant development. Although there are many plant hormones, six have been well-characterized: auxins, gibberellins, cytokinins, ethylene, abscisic acid, and brassinosteroids.

38.3 Plants have different receptors that detect various colors of light. Phytochrome detects red and far-red light, which affect several aspects of development, including the timing of flowering.

he ultimate control of plant growth and development, which includes all the changes that take place during the entire life of an individual,

are not present, that characteristic does not develop. When a particular gene is present, its expression—that is, how it exhibits itself as an observable feature of an organism—is determined by several factors, including signals from other genes and from the environment. The location of a cell in the young plant body also has a profound effect on gene expression during development. Chemical signals from adjacent cells help a cell “perceive” its location within the plant body. Each cell’s spatial environment helps determine what that cell ultimately becomes. Growth and development, including a plant’s responses to various changes in its environment, are controlled by plant hormones, organic

38.4 The exposure of a plant to a herbivore or pathogen

molecules that are present in very low concentrations in plant tissues and

causes a localized hypersensitive response that seals off the affected area. The plant also sends signals from the site of localized infection, leading to systemic acquired resistance—long-lasting, enhanced defenses throughout the plant.

that act as chemical signals between cells. Environmental cues, such as changing day length and variations in precipitation and temperature, exert an important influence on gene expression and hormone production, as they do on all aspects of plant growth and development.

For example, the initiation of sexual reproduction is often under environmental control, particularly in temperate latitudes, and plants switch from vegetative to reproductive growth after receiving appropriate signals from the environment. Many flowering plants are sensitive to changes in the relative amounts of daylight and darkness that accompany the changing seasons, and these plants flower in response to those changes (see photograph). In this chapter we will see that plants use light as an external signal © Martin Shields/Alamy

to help regulate many aspects of growth and development, such as tropisms (directional growth), time of flowering, shade avoidance, and circadian rhythms. Other plants have temperature requirements that induce sexual reproduction. Thus, plants continually perceive information from the environment and use this information to help regulate normal growth and development. We consider these various aspects of growth and development in this chapter.

38.1 TROPISMS ■ ■ LEARNING OBJECTIVE Describe phototropism, gravitropism, and thigmotropism.

Plants exhibit movements in response to environmental stimuli such as light, gravity, and touch. A plant may respond to such an external stimulus by directional growth—that is, the direction of growth depends on the direction of the stimulus. Such a directional growth response, called a tropism, results in a change in the position of a plant part. Tropisms are irreversible and may be positive or negative, depending on whether the plant grows toward the stimulus (a positive tropism) or away from it (a negative tropism). Tropisms are under hormonal control, which is discussed later in the chapter. Phototropism is the directional growth of a plant caused by light (FIG. 38-1). Most growing shoot tips exhibit positive phototropism by bending (growing) toward light, something you may have observed if you place houseplants near a sunny window. This growth response

On day 3, turned on its side

increases the likelihood that stems and leaves receive adequate light for photosynthesis. The bending response of phototropism is triggered by blue light with wavelengths less than 500 nm. (You may recall from Chapter 34 that blue light also induces stomata to open.) For a plant or any organism to have a biological response to light, it must contain a light-sensitive substance, called a photoreceptor, to absorb the light. The photoreceptor that absorbs blue light and triggers the phototropic response and other blue-light responses (such as stomatal opening) is a family of yellow pigments called phototropins. Phototropins are light-activated kinases, enzymes that transfer phosphate groups. There is evidence that phototropins become phosphorylated—that is, a phosphate group is added—in response to blue light. Thus, phosphorylation is an early step in the blue-light-signaling pathway. Growth in response to the direction of gravity is called gravitropism. Most stem tips exhibit negative gravitropism by growing away from Earth’s center, whereas most root tips exhibit positive gravitropism (FIG. 38-2). The root cap is the site of gravity perception in

One hour later

(a)

Cengage

1

FIGURE 38-1 Phototropism A bean seedling (Phaseolus vulgaris) grows in the direction of light and therefore exhibits positive phototropism. The bending is caused by greater elongation on the shaded side of the stem than on the lighted side.

FIGURE 38-2 Animated Gravitropism (a) A corn (Zea mays) seedling was turned on its side three days after germinating. In 1 hour the root and shoot tips had curved. (b) In 24 hours (seedling on the right), the new root growth was downward (positive gravitropism) and the new shoot growth was upward (negative gravitropism). A control seedling germinated at the same time is on the left.

(b)

roots; when the root cap is removed, the root continues to grow, but it loses any ability to perceive gravity. Special cells in the root cap possess starch-containing amyloplasts that collect toward the bottom of the cells in response to gravity, and these amyloplasts may initiate at least some of the gravitropic response. If the root is put in a different position, as when a potted plant is laid on its side, the amyloplasts tumble to a new position, always settling in the direction of gravity. The gravitropic response (bending) occurs shortly thereafter and involves the hormone auxin (discussed later in the chapter). Despite the movement of amyloplasts in response to gravity, researchers question their role in gravitropism. A mutant Arabidopsis thaliana plant that lacks amyloplasts in its root cap cells still responds gravitropically when placed on its side, indicating that roots do not necessarily need amyloplasts to respond to gravity. Ongoing research may clarify how roots perceive gravity. Thigmotropism is growth in response to a mechanical stimulus, such as contact with a solid object. The twining or curling growth of tendrils or stems, which helps attach a climbing plant such as a vine to some type of support, is an example of thigmotropism (see Fig. 34-14b).

Review ■

■

What is phototropism? How does blue light trigger the phototropic response? How do phototropism and gravitropism differ?

38.2 PLANT HORMONES AND DEVELOPMENT ■ ■ LEARNING OBJECTIVES 2 3 4

Describe a general mechanism of action for plant hormones, using auxin as your example. Describe early auxin experiments involving phototropism. List several ways that each of these hormones affects plant growth and development: auxins, gibberellins, cytokinins, ethylene, abscisic acid, and brassinosteroids.

A plant hormone is an organic compound that acts as a chemical signal eliciting a variety of responses that regulate growth and

TABLE 38-1

development. The study of plant hormones is challenging because hormones are effective in extremely small concentrations (less than 10−6 mol/L). In addition, the effects of different plant hormones overlap, and it is difficult to determine which hormone, if any, is the primary cause of a particular response. Plant hormones may also stimulate a response at one concentration and inhibit that same response at a different concentration. For many years, biologists studied five major classes of plant hormones: auxins, gibberellins, cytokinins, ethylene, and abscisic acid. More recently, researchers have uncovered compelling evidence for brassinosteroids as a sixth plant hormone. Additional plant hormones will probably be characterized in the future. (TABLE 38-1 summarizes the six plant hormones.)

Plant hormones act by signal transduction Researchers have used molecular genetic techniques to better understand the biology of plant hormones. Arabidopsis mutants are particularly useful. For example, different mutants have defects in hormone synthesis, hormone transport, signal reception, or signal transduction (FIG. 38-3). In signal transduction, a receptor converts an extracellular signal into an intracellular signal that causes one or more cellular responses. Such mutants enable plant biologists to identify and clone genes involved in these aspects of hormone biology. Studying the mutant phenotypes helps plant biologists establish connections between the mutant genes and specific physiological activities involved in growth and development. Using this research, biologists are elucidating the general mechanisms of plant hormone action, an important goal of systems biology. It appears that many plant hormones bind to enzyme-linked receptors; the hormone binds to the receptor, where it triggers an enzymatic reaction of some sort. Let us consider a specific example that involves the plant hormone auxin, which is known to cause rapid changes in gene expression. One receptor for auxin is located in either the cytosol or the nucleus and has a three-dimensional shape that binds to the auxin molecule. The binding of auxin to its receptor (designated the TIR1 receptor) catalyzes the attachment of the molecule ubiquitin to repressor proteins that inhibit the transcription of certain genes, designated auxin response genes (FIG. 38-4).

Plant Hormones

Hormone

Site of Production

Principal Actions

Auxins (e.g., IAA) Gibberellins (e.g., GA3) Cytokinins (e.g., zeatin) Ethylene

Shoot apical meristem, young leaves, seeds Young leaves and shoot apical meristems, embryo in seed Various sites in plants

Stem elongation, apical dominance, root initiation, fruit development Seed germination, stem elongation, flowering, fruit development

Abscisic acid Brassinosteroids (e.g., brassinolide)

Stem nodes, ripening fruit, damaged or senescing tissue Almost all cells that contain plastids (leaves, stems, roots) Shoots (leaves and flower buds), seeds, fruits

Cell division, delay of leaf senescence, inhibition of apical dominance, flower development, embryo development, seed germination Fruit ripening, responses to environmental stressors, seed germination, maintenance of apical hook on seedlings, root initiation, senescence and abscission in leaves and flowers Seed dormancy, responses to water stress Light-mediated gene expression, cell division, cell elongation, seed germination, vascular development

External (environmental) signals Auxin synthesis

Directional (polar) auxin transport

Reception, signal transduction, and response

Local auxin gradient

Specific developmental responses

Internal signals

FIGURE 38-3 General mechanism of action for the hormone auxin Various external and/or internal signals trigger the synthesis of auxin. Once synthesized, auxin is transported directionally, causing local auxin gradients to form in the plant. In turn, these gradients trigger specific developmental responses in certain cells by reception, signal transduction, and response (discussed shortly). Arabidopsis mutants have been identified with defects involving auxin synthesis, auxin transport, or other steps in the process.

Enzyme complex Auxin receptor (TlR1)

Repressor protein (inhibits transcription) Transcription activator Auxin response element

DNA No transcription

Auxin response gene

(a) Auxin absent. A repressor protein binds to a transcription activator, inhibiting transcription of the auxin response gene.

Proteasome Enzyme complex

Ubiquitinylated protein

Peptide fragments

Auxin Ubiquitin molecules Transcription activator DNA Transcription occurs

Auxin response gene

(b) Auxin present. Auxin binds to the TlR1 receptor, which is part of an enzyme complex that attaches ubiquitin to the repressor protein. The ubiquitinylated rerpressor protein is then degraded into peptide fragments in a proteasome. With the repressor protein no longer bound to the transcription activator, transcription of the auxin response gene occurs.

FIGURE 38-4 Model of a signal transduction pathway for auxin

The ubiquitinylated protein is therefore targeted for degradation into peptide fragments in a proteasome. (See Figure 14-15, which shows the ubiquitin-proteasome degradation of proteins.) As a result, transcription of the auxin response gene occurs, resulting in changes in cell growth and development.

The identification of the TIR1 receptor for auxin was reported in 2005 by two groups working independently. This receptor is an F-box protein. The F-box is a short sequence of amino acids found in molecules that catalyze the addition of ubiquitin tags to proteins targeted for destruction. Interestingly, all animals, plants,

and fungi examined to date use ubiquitin to target proteins for destruction, but bacteria do not. This suggests that the use of ubiquitin tags evolved early in eukaryote evolution and was conserved as the various groups of eukaryotes diversified. Plants have about 700 different F-box proteins. Although not much is known about most of them, it is significant that researchers have identified other plant hormones and signaling molecules that use F-box proteins as receptors. These include gibberellin, ethylene, and abscisic acid.

Auxins promote cell elongation Charles Darwin, the British naturalist best known for developing the theory of natural selection to explain evolution, also provided the first evidence for the existence of auxins. The experiments that Darwin and his son Francis performed in the 1870s involved positive phototropism, the directional growth of plants toward light. The plants they used were newly germinated canary grass seedlings. As in all grasses, the first part of a canary grass seedling to emerge from the soil is the coleoptile, a protective sheath that encircles the stem. When coleoptiles are exposed to light from only one direction, they bend toward the light. The bending occurs below the tip of the coleoptile. The Darwins tried to influence this bending in several ways (FIG. 38-5). For example, they covered the tip of the coleoptile as soon as it emerged from the soil. When they covered that part of the coleoptile above where the bend would be expected to occur, the plants did not bend. On other plants, they removed the coleoptile tip and found bending did not occur. When the bottom of the coleoptile where the curvature would occur was shielded from the

light, the coleoptile bent toward light. From these experiments, the Darwins concluded that “some influence is transmitted from the upper to the lower part, causing it to bend.” In the 1920s, Frits Went, a young Dutch scientist, isolated the phototropic hormone from oat coleoptiles. He removed the coleoptile tips and placed them on tiny blocks of agar for a period of time. When he put one of these agar blocks squarely on a decapitated coleoptile, normal growth resumed. When he placed one of these agar blocks to one side of the tip of a decapitated coleoptile in the dark, bending occurred (FIG. 38-6). The results indicated that the substance had diffused from the coleoptile tip into the agar and, later, from the agar into the decapitated coleoptile. Went named this substance auxin (from the Greek aux, “enlarge” or “increase”). Auxin is a group of several natural (and artificial) plant hormones; the most common and physiologically important auxin is indoleacetic acid (IAA). The purification and elucidation of the primary auxin’s chemical structure were accomplished in the mid1930s by a research team led by Kenneth Thimann at the California Institute of Technology. The movement of auxin in the plant is said to be polar, or unidirectional. Auxin moves downward along the shoot–root axis from its site of production, usually the shoot apical meristem. (Young leaves and seeds are also sites of auxin production.) This directional cell to cell transport results in an auxin gradient (a spatial difference in auxin concentration) that triggers specific responses. Special auxin transport proteins in the plasma membranes of plant cells control the movement of auxin out of cells; this process is responsible for the directional movement of auxin. Mutations in

KEY EXPERIMENT QUESTION: What part of the grass coleoptile depends on light for phototropic growth? HYPOTHESIS: The coleoptile tip is required for the growth (bending) of the coleoptile shaft that accompanies exposure to light coming from one direction.

EXPERIMENT: Some canary grass coleoptiles were uncovered, some were covered only at the tip, some had the tip removed, and some were covered everywhere but at the tip (left). The covers were impervious to light.

Light rays

RESULTS AND CONCLUSION: After exposure to light coming from one direction, the uncovered plants and the plants with uncovered tips grew toward the light (right). The plants with tips covered or removed did not bend toward light. The Darwins concluded that some substance was produced in the tip and transmitted to the lower part, causing it to bend. Sources: Darwin, C. R. The Power of Movement in Plants. John Murray, London, 1880; Boysen-Jensen, P. “Concerning the performance of phototropic stimuli on the Avena coleoptile.” Berichte der Deutschen Botanischen Gesellschaft (Reports of the German Botanical Society), Vol. 31, pp. 559–566, 1913.

FIGURE 38-5 The Darwins’ phototropism experiments

Shaded side of coleoptile

KEY EXPERIMENT QUESTION: Is there a chemical substance responsible for elongation of coleoptiles, and can it be isolated? HYPOTHESIS: The factor responsible for coleoptile growth is a diffusible chemical that can be isolated from coleoptile tips. EXPERIMENT: Coleoptile tips were placed on agar blocks for a period of time (a). The agar block was transferred to a decapitated coleoptile. It was placed off-center, and the coleoptile was left in darkness (b). Light rays

Coleoptile tip

Illuminated side of coleoptile

(a)

Agar block

(b)

(c)

FIGURE 38-7 Animated Phototropism and the unequal distribution of auxin RESULTS AND CONCLUSION: The coleoptile bent (c), indicating that a chemical moved from the original coleoptile tip to the agar block and from there to the same side of the decapitated coleoptile, causing that side to elongate. Source: Went, F. “A growth substance and growth.” Recueils Travaux Botaniques Néerlandais (Collections of Dutch Botanical Works), Vol. 25, pp. 1–116, 1928.

FIGURE 38-6 Animated Isolating auxin from coleoptiles

auxin transport proteins inhibit the transmission of auxin, which normally moves about 1 cm per hour. Auxin’s most characteristic action is promotion of cell elongation in stems and coleoptiles. This effect, apparently exerted by acidification of cell walls, increases their plasticity and thus enables them to expand under the force of the cell’s internal turgor pressure. Auxin’s effect on cell elongation also provides an explanation for phototropism. When a plant is exposed to a light from only one direction, some of the auxin migrates laterally to the shaded side of the stem before moving down the stem by polar transport. Because of the greater auxin concentration on the shaded side of the stem, the cells there elongate more than the cells on the light side, and the stem bends toward the light (FIG. 38-7). Auxin is also involved in gravitropism and thigmotropism. Auxin exerts other effects on plants. For example, some plants tend to branch out very little when they grow. Growth in these plants occurs almost exclusively from the apical meristem rather than from axillary buds, which do not develop as long as the terminal bud is present. Such plants are said to exhibit apical dominance, the inhibition of axillary bud growth by the apical meristem. In plants with strong apical dominance, auxin produced

Auxin travels down the side of the stem or coleoptile away from the light, causing cells on the shaded side to elongate. Therefore, the stem or coleoptile bends toward light.

in the apical meristem inhibits axillary buds near the apical meristem from developing into actively growing shoots. When the apical meristem is pinched off, the auxin source is removed and axillary buds grow to form branches. Apical dominance is often quickly re-established, however, as one branch begins to inhibit the growth of others. Other hormones (ethylene and cytokinin, both discussed later) are also involved in apical dominance. As with many physiological activities, the changing ratios of these hormones may be the factor responsible for apical dominance. Auxin produced by developing seeds stimulates the development of the fruit. When auxin is applied to certain flowers in which fertilization has not occurred (and, therefore, in which seeds are not developing), the ovary enlarges and develops into a seedless fruit. Seedless tomatoes are produced in this manner.1 Auxin is not the only hormone involved in fruit development, however. Some manufactured, or synthetic, auxins have structures similar to that of IAA. The synthetic auxin naphthalene acetic acid stimulates root development on stem cuttings and is used for asexual propagation, particularly of woody plants with horticultural importance (FIG. 38-8). The synthetic auxins 2,4-D and 2,4,5-T have been used as selective herbicides (weed killers). These compounds kill plants with broad leaves but, for reasons not completely under1 Not all seedless fruits are produced by treatment with auxin. In Thompson seedless grapes, fertilizations occurs but the embryos abort, and therefore the seeds fail to develop. Thompson seedless grapes are sprayed with the hormone gibberellin to increase berry size.

Joe Eakes, Color Advantage/Visuals Unlimited

Robert Lyons/Visuals Unlimited

FIGURE 38-8 Auxin and root development on stem cuttings Many adventitious roots developed on a honeysuckle (Lonicera fragrantissima) cutting placed in a solution with a high concentration of synthetic auxin (left). Fewer roots developed in a lower auxin concentration (middle). The cutting placed in water (no auxin) served as a control and did not form roots in the same period (right).

stood, do not kill grasses. Both herbicides are similar in structure to IAA and disrupt the plants’ normal growth processes. Because many of the world’s most important crops are grasses (such as wheat, corn, and rice), 2,4-D and 2,4,5-T can kill broadleaf weeds that compete with these crops. The use of 2,4,5-T is no longer allowed in the United States, however, because of its association with dioxins, a group of mildly to very toxic compounds formed as byproducts when 2,4,5-T is manufactured.

Gibberellins promote stem elongation In the 1920s and 1930s, a Japanese biologist was studying a disease of rice in which the young rice seedlings grew extremely tall and spindly, fell over, and died. The cause of the disease was a fungus (Gibberella fujikuroi) that produces a chemical substance named gibberellin (GA). Not until after World War II did scientists in Europe and North America learn of the work done by the Japanese scientists. During the 1950s and 1960s, studies in the United States and Great Britain showed that gibberellins are produced by healthy plants as well as by the fungus. Gibberellins are hormones involved in many normal plant functions. The symptoms of the seedling disease are caused by an abnormally high GA concentration in the plant tissue (because both plant and fungus produce gibberellin). Currently, dozens of naturally occurring GAs are known, although only about four function as plant hormones; the rest are inactive precursors. There are no synthetic gibberellins. Gibberellins promote stem elongation in many plants. When GA is applied to a plant, particularly certain dwarf varieties, this elongation may be spectacular. Some corn and pea plants that are dwarfs as a result of one or more mutations grow to a normal height when treated with GA. Short-stemmed, high-yielding varieties of wheat have short stems because they have a reduced response to GA. These varieties put less of their resources into stem height and more resources into grain production. Gibberellins are also involved in bolting, the rapid elongation of a floral stalk that occurs naturally in many plants when they initiate flowering (FIG. 38-9).

FIGURE 38-9 Gibberellin and stem elongation Like many biennials, Indian blanket (Gaillardia pulchella) grows as a rosette, which is a circular cluster of leaves close to the ground, during its first year (left). It then bolts when it initiates flowering in the second year (right). The plant in bloom grows to 0.6 m (2 ft).

Gibberellins cause stem elongation by stimulating cells to divide as well as to elongate. The actual mechanism of cell elongation differs from that caused by auxin, however. Recall that IAAinduced cell elongation involves the acidification of the cell wall. In GA-induced cell elongation, cell wall acidification does not occur; biologists do not yet know how gibberellins modify cell wall properties to cause cell elongation. Gibberellins affect several reproductive processes in plants. They stimulate flowering, particularly in long-day plants (discussed later in the chapter). In addition, GAs substitute for the low temperature that biennials require before they begin flowering. If GAs are applied to biennials during their first year of growth, flowering occurs without exposure to a period of low temperature. Gibberellins, like auxin, affect fruit development. Agriculturalists apply GAs to several varieties of grapes to produce larger fruits. Gibberellins are also involved in seed germination in certain plants. In a classic experiment involving barley seed germination, researchers showed that the release of GA from the embryo triggers the synthesis of α-amylase, an enzyme that digests starch in the endosperm. As a result, glucose becomes available for absorption by the embryo. Although enzymes mobilize starch reserves in many types of seeds, GA control of seed enzymes appears restricted to cereals and other grasses. In addition to mobilizing food reserves in newly germinated grass seeds, application of gibberellins substitutes for low-temperature or light requirements for germination in seeds of plants such as lettuce, oats, and tobacco. Scientists are actively conducting research in gibberellin signaling and have identified several GA receptors in the nucleus, repressor proteins designated DELLA proteins, as well as an F-box protein. When GA is not bound to the receptor, a DELLA protein

Cell division without differentiation

inhibits transcription of GA response genes. When GA is present, it binds to the receptor, which triggers an enzyme that attaches ubiquitin to the DELLA protein. This protein is targeted for degradation into peptide fragments in a proteasome. With the DELLA protein no longer present, transcription of the GA response gene occurs. This signaling process may sound familiar: the GA signal transduction process shares similarities with the auxin signal transduction pathway described earlier in the chapter.

Cell division with differentiation

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During the 1940s and 1950s, researchers were trying to find substances that might induce plant cells to divide in tissue culture, a technique in which cells are isolated from plants and grown in a nutrient medium. (Plant tissue culture is discussed in Chapter 33.) They discovered that cells would not divide without a substance found in coconut milk. Because coconut milk has a complex chemical composition, investigators did not chemically identify the division-inducing substance for some time. Finally, researchers isolated an active substance from a different source, aged DNA from herring sperm. They called it cytokinin because it induces cytokinesis, or cytoplasmic division. In 1963, researchers identified the first plant cytokinin, zeatin, in corn. Since then, similar molecules have been identified in other plants. Biologists have also synthesized several cytokinins. Cytokinins are structurally similar to adenine, a purine base that is part of DNA and RNA molecules. Biologists are beginning to elucidate the cytokinin signal transduction pathway, which is more complex than those of auxin and gibberellin. The cytokinin pathway has multiple steps, each under the control of a number of genes. The presence of feedback loops and links to other signaling pathways adds another level of complexity to the regulation of cytokinin activity. Cytokinins promote cell division and differentiation of young, relatively unspecialized cells into mature, more specialized cells in intact plants. They are a required ingredient in any plant tissue culture medium and must be present for cells to divide. In tissue culture, cytokinins interact with auxin during the formation of plant organs such as roots and stems (FIG. 38-10). For example, in tobacco tissue culture a high ratio of auxin to cytokinin induces root formation, whereas a low ratio of auxin to cytokinin induces shoot formation. Cytokinins and auxin also interact in the control of apical dominance. Here their relationship is antagonistic: auxin inhibits the growth of axillary buds, and cytokinin promotes their growth. One effect of cytokinins on plant cells is to delay the aging process. Plant cells, like all living cells, go through a natural aging process known as senescence. Senescence is accelerated in cells of plant parts that are cut, such as flower stems. Botanists think that plants must have a continual supply of cytokinins from the roots. Cut stems, of course, lose their source of cytokinins and therefore age rapidly. When cytokinins are sprayed on leaves of a cut stem of many species, they remain green, whereas unsprayed leaves turn yellow and die. Despite their involvement in regulating many aspects of plant growth and development, cytokinins currently have few commercial applications other than plant tissue culture. However, in 1995,

Cytokinin

FIGURE 38-10 Auxin–cytokinin interactions in tissue culture Varying amounts of auxin and cytokinin in the culture media produce different growth responses. (a) The initial explant is a small piece of sterile tissue from the pith of a tobacco stem, which is placed on nutrient agar. (b) Nutrient agar containing 2.0 mg/L of auxin and 0.2 mg/L of cytokinin causes cells to divide and form a clump of undifferentiated tissue called a callus. (c) Agar with 2.0 mg/L of auxin and 0.02 mg/L of cytokinin (high ratio of auxin to cytokinin) stimulates root growth. (d) Agar with 2.0 mg/L of auxin and 0.5 mg/L of cytokinin (low ratio of auxin to cytokinin) stimulates shoot growth.

molecular biologists at the University of Wisconsin combined a promoter from a gene activated during normal senescence with a gene that encodes an enzyme involved in cytokinin synthesis. (Recall from Chapter 13 that the promoter is the nucleotide sequence in DNA to which RNA polymerase attaches to begin transcription.) The leaves of transgenic tobacco plants that contained this recombinant DNA produced more cytokinin and therefore lived longer and continued to photosynthesize (FIG. 38-11). This molecular technology may have the potential to increase the longevity and productivity of certain crops.

Ethylene promotes abscission and fruit ripening During the early 20th century, scientists observed that the gas ethylene (C2H4) has several effects on plant growth, but not until 1934 did they demonstrate that plants produce ethylene. This natural hormone influences many plant processes. Ethylene inhibits cell elongation, promotes seed germination, promotes apical dominance, and is involved in plant responses to wounding or invasion by disease-causing microorganisms. Ethylene also has a major role in many aspects of senescence, including fruit ripening. As a fruit ripens, it produces ethylene, which triggers an acceleration of the ripening process. This induces

Ethylene is involved in leaf abscission, which is actually influenced by two antagonistic hormones, ethylene and auxin. As a leaf ages (as autumn approaches, for deciduous trees in temperate climates), the level of auxin in the leaf decreases. Concurrently, cells in the abscission layer at the base of the petiole (where the leaf will break away from the stem) begin producing ethylene. Researchers studying ethylene-signaling have identified a similar mechanism to that already described for auxin and gibberellin. When ethylene is present, it binds to a receptor. This binding triggers an enzyme that attaches ubiquitin to a protein targeted for degradation into peptide fragments in a proteasome. At least two F-box proteins are involved in the protein degradation that occurs in ethylene signal transduction.

Courtesy of Dr. Richard M. Amasino, University of Wisconsin

Abscisic acid promotes seed dormancy

FIGURE 38-11 Cytokinin synthesis and delay of senescence

the fruit to produce more ethylene, which further accelerates ripening. The expression “one rotten apple spoils the barrel” is true. A rotten apple is one that is overripe and produces large amounts of ethylene, which diffuses and triggers the ripening process in nearby apples. Humans use ethylene commercially to uniformly ripen bananas and tomatoes. These fruits are picked while green and shipped to their destination, where they are exposed to ethylene before they are delivered to grocery stores (FIG. 38-12). Plants growing in a natural environment encounter rain, hail, wind, and contact with passing animals. All these mechanical stressors alter their growth and development, making them shorter and stockier than plants grown in a greenhouse. Ethylene regulates such developmental responses to mechanical stimuli, known as thigmomorphogenesis. Plants that are mechanically disturbed produce additional ethylene, which in turn inhibits stem elongation and enhances cell wall thickening in supporting cells (collenchyma and sclerenchyma). These changes are adaptive because shorter, thicker stems are less likely to be damaged by mechanical stressors.

Illustration Services

The tobacco (Nicotiana tabacum) plant on the left was genetically engineered to produce additional cytokinin as it aged, whereas the tobacco plant of the same age on the right served as a control. Note the extensive senescence and death of older leaves on the control plant. Depending on the variety, tobacco grows 0.9 to 3 m (3 to 10 ft) tall.

In 1963, two different research teams discovered abscisic acid (ABA). Despite its name, abscisic acid does not induce abscission in most plants. Instead, ABA is involved in a plant’s response to stress and in seed dormancy, a temporary state of arrested physiological activity. As an environmental stress hormone, ABA particularly promotes changes in plant tissues that are water-stressed. (Recall that ethylene also affects plant responses to certain stressors, such as mechanical stressors and wounding.) Botanists best understand the effect of abscisic acid on plants suffering from water stress. The level of ABA increases dramatically in the leaves of plants exposed to severe drought conditions. The high level of ABA in the leaves activates a signal transduction process that leads to the closing of stomata. The closing of stomata saves the water that the plant would normally lose by transpiration, thereby increasing the plant’s likelihood of survival. As knowledge of ABA signaling in guard cells increases, botanists hope to use this information to engineer crops and horticultural plants that are resistant to drought.

FIGURE 38-12 Ethylene and fruit ripening Both boxes of tomatoes were picked at the same time, while green. The tomatoes in the box on the right were exposed to an atmosphere containing 100 ppm of ethylene for three days.

Courtesy of M. G. Neuffer

verse this defect by applying BR exogenously to the plant. Studies of these mutants suggest that brassinosteroids are involved in multiple developmental processes, such as cell division, cell elongation, light-induced differentiation, vascular development, and seed germination (FIG. 38-14). Many steps of the BR signaling pathway have been characterized. BRs bind to a receptor kinase in the plant plasma membrane (unlike steroid hormones in animals, which enter the cell and bind to receptors in the cytoplasm or nucleus). When BR binds to its receptor, the receptor phosphorylates molecules in the cytosol, which in turn triggers a signaling cascade that ultimately alters gene expression in the cell. Part of the signal transduction process involves attaching ubiquitin to nuclear proteins targeted for degradation in a proteasome. As a result, the expression of BR target genes is adjusted.

Identification of a universal flower-promoting signal remains elusive Experiments in which different tobacco species are grafted together indicate that unidentified flower-promoting and flower-inhibiting

FIGURE 38-13 The absence of abscisic acid causes precocious germination

100

In a corn (Zea mays) mutant that does not produce abscisic acid, some of the kernels have germinated while still on the ear, producing roots (arrows).

Brassinosteroids are plant steroid hormones Although steroid hormones have long been known to have crucial roles in animals, the roles of steroid hormones in plants were not well characterized until the mid-1990s. The brassinosteroids (BRs) are a group of steroids that function as plant hormones. BRs are involved in several aspects of growth and development. Arabidopsis mutants that cannot synthesize BRs are dwarf plants with small curly leaves and reduced fertility; researchers can re-

Percentage germination

The onset of winter is also a type of stress on plants. A winter adaptation that involves abscisic acid is dormancy in seeds. Many seeds have high levels of ABA in their tissues and do not germinate until the ABA washes out. In a corn mutant unable to synthesize ABA, the seeds germinate as soon as the embryos are mature, even while attached to the ear (FIG. 38-13). Abscisic acid is not the only hormone involved in seed dormancy. For example, addition of gibberellin reverses the effects of dormancy. In seeds, the level of ABA decreases during the winter, and the level of GA increases. Cytokinins are also implicated in breaking dormancy. Once again, you see that a single physiological activity such as seed dormancy may be controlled in plants by the interaction of several hormones. The plant’s actual response may result from changing ratios of hormones rather than the effect of each individual hormone.

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FIGURE 38-14 Brassinosteroids and seed germination in Arabidopsis Researchers used a GA-insensitive Arabidopsis mutant that produced seeds that did not germinate. (Recall that GA promotes seed germination, so a plant that does not respond to endogenous GA is required.) The scientists overcame the inability of the mutant seeds to germinate by treating them with a brassinosteroid (BR). Note how the percentage of germination increases as the concentration of BR to which the seeds were exposed increases. Original source: Steber, C. M. and P. McCourt. “A role for brassinosteroids in germination in Arabidopsis.” Plant Physiology, Vol. 125, pp. 287–297, 2001.

38.3 LIGHT SIGNALS AND PLANT DEVELOPMENT

Graft

Day-neutral plant grafted to long-day plant

■ ■ LEARNING OBJECTIVES

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sion of phytochrome signal transduction.

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FIGURE 38-15 Evidence for the existence of a flower-promoting substance When a long-day tobacco plant (Nicotiana silvestris) is grafted to a day-neutral tobacco plant (N. tabacum) and both plants are exposed to a long-day/ short-night regimen, they both flower. The day-neutral plant flowers sooner than it normally would, presumably because a flower-promoting substance passes from the long-day plant to the day-neutral one through the graft.

substances may exist. Nicotiana silvestris is a long-day tobacco plant (it requires a short night to flower); a variety of N. tabacum is a dayneutral tobacco plant (seasonal changes do not affect when it flowers). When a long-day tobacco is grafted to a day-neutral tobacco and exposed to short nights, both plants flower (FIG. 38-15). The day-neutral tobacco plant flowers sooner than it normally would. Biologists hypothesize that a flower-promoting substance, florigen, may be induced in the long-day plant and transported to the day-neutral plant through the graft union, causing the day-neutral tobacco plant to flower sooner than expected. An intact plant may produce florigen in the leaves and transport it in the phloem to the shoot apical meristem. There, it induces a transition from vegetative to reproductive development—that is, to a meristem that produces flowers. When a botanist grafts a long-day tobacco to a day-neutral tobacco and exposes them to long nights, neither plant flowers. As long as these conditions continue, the day-neutral plants do not flower even when they would normally do so. In this case, the long-day tobacco may produce a flower-inhibiting substance that is transported to the day-neutral tobacco through the graft union. This substance prevents the day-neutral tobacco from flowering.

Review ■

■ ■

Explain how varying amounts of light and darkness induce flowering.

6 Describe the role of phytochrome in flowering, including a brief discus-

How are signal reception, signal transduction, and altered cell activity part of the general mechanism of action for auxin? How is auxin involved in phototropism? Which hormones are involved in each of the following physiological processes: (1) seed germination, (2) stem elongation, (3) fruit ripening, (4) leaf abscission, and (5) seed dormancy?

7

Define circadian rhythm and give an example.

8

Distinguish between phytochrome and cryptochrome.

Photoperiodism is any response of a plant to the relative lengths of daylight and darkness. Initiation of flowering at the shoot apical meristem is one of several physiological activities that are photoperiodic in many plants. Plants are classified into four main groups— short-day, long-day, intermediate-day, and day-neutral—on the basis of how photoperiodism affects their transition from vegetative growth to flowering. Short-day plants (also called long-night plants) flower when the night length is equal to or greater than some critical period (FIG. 38-16a). Thus, short-day plants detect the lengthening nights of late summer or fall. There are two kinds of short-day plants: qualitative and quantitative. In qualitative short-day plants, flowering occurs only in short days; in quantitative short-day plants, flowering is accelerated by short days. The initiation of flowering in short-day plants is not due to the shorter period of daylight but to the long, uninterrupted period of darkness. The minimum critical night length varies considerably from one plant species to another but falls between 12 and 14 hours for many. Examples of shortday plants are florist’s chrysanthemum, cocklebur, and poinsettia, which typically flower in late summer or fall. Poinsettias, for example, typically initiate flower buds in early October in the Northern Hemisphere and flower about 8 to 10 weeks later; hence, their traditional association with Christmas. Long-day plants (also called short-night plants) flower when the night length is equal to or less than some critical period (FIG. 38-16b). In qualitative long-day plants, flowering occurs only in long days; in quantitative long-day plants, long days accelerate flowering. Long-day plants, such as spinach, black-eyed Susan (see photograph in chapter introduction), and the model research plant Arabidopsis, flower when they detect the shortening nights of spring and early summer. Intermediate-day plants do not flower when night length is either too long or too short (FIG. 38-16c). Sugarcane and coleus are intermediate-day plants. These plants flower when exposed to days and nights of intermediate length. Some plants, called day-neutral plants, do not initiate flowering in response to seasonal changes in the period of daylight and darkness but instead respond to some other type of stimulus, external or internal. Cucumber, sunflower, corn, and onion are examples of day-neutral plants. Many of these plants originated in the tropics, where day length does not vary appreciably during the year. In contrast, short-day, long-day, and intermediate-day plants are temperate or subtropical (midlatitude species).

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(b) Long-day plants flower when the night length is equal to or less than a certain critical length in a 24-hour period. The critical length varies among different species; short-day plants may require a shorter night length than long-day plants, as shown.

FIGURE 38-17 Animated Photoperiodic responses of short-day and long-day plants A short-day plant flowers when it is grown under long-night conditions (top middle), but it does not flower when exposed to a long night interrupted with a brief flash of red light (top right). A long-day plant does not flower when grown under long-night conditions (bottom middle) unless the long night is interrupted with a brief flash of red light (bottom right).

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FIGURE 38-16 Generalized photoperiodic responses in short-day, long-day, and intermediate-day plants

Plant biologists have experimented with the effects of various light regimens on flowering. FIGURE 38-17 shows how flowering in long-day and short-day plants is affected by different light treatments, including a short-day/long-night regimen with a night break, a short burst of light in the middle of the night.

Phytochrome detects day length The main photoreceptor for photoperiodism and many other light-initiated plant responses (such as germination and seedling establishment) is phytochrome, a family of about five blue-green pigment proteins, each of which is coded for by a different gene. A mixture of phytochrome proteins is present in cells of all vascular plants examined so far. For example, five members of the phytochrome family, designated phyA, phyB, phyC, phyD, and phyE, oc-

cur in Arabidopsis. Phytochrome is also a photoreceptor in certain bacteria, fungi, and slime molds. Much of the current knowledge of phytochrome in Arabidopsis is based on various mutant plants that do not express a specific phytochrome gene, such as the gene that codes for phyA. By studying the physiological response of plants that do not produce an individual phytochrome, biologists have concluded that the individual forms of phytochrome have both unique and overlapping functions. PhyB appears to exert its influence at all stages of the plant life cycle, whereas the other forms of phytochrome have narrower functions at specific stages in the life cycle. PhyA and phyB may have antagonistic (opposite) effects on flowering. In long-day plants, phyB may inhibit flowering and phyA may induce flowering. Flowering occurs more rapidly in long-day plants with mutations in the gene that codes for phyB; such mutations reduce or eliminate the production of phyB. In contrast, flowering is delayed or prevented in long-day plants with mutations in the gene that codes for phyA. Each member of the phytochrome family exists in two forms and readily converts from one form to the other after absorption of light of specific wavelengths. One form, designated Pr (for redabsorbing phytochrome), strongly absorbs light with a relatively short red wavelength (660 nm). In the process, the shape of the molecule changes to the second form of phytochrome, Pfr, so

Phytochrome synthesis

Phytochrome degradation by a proteasome

Red light (660 nm) Regulation of gene expression Far-red light Pr

(730 nm)

Physiological response (flowering, seed germination, etc.)

Pfr

FIGURE 38-18 Animated Phytochrome Each member of the phytochrome family occurs in two forms, designated Pr and Pfr, and readily converts from one form to the other. Red light (660 nm) converts Pr to Pfr, and far-red light (730 nm) converts Pfr to Pr.

designated because it absorbs far-red light, which is light with a relatively long red wavelength (730 nm; FIG. 38-18). When it absorbs far-red light, Pfr reverts to the original form, Pr. Pfr is the active form of phytochrome, triggering or inhibiting physiological responses such as flowering. What does a pigment that absorbs red light and far-red light have to do with daylight and darkness? Sunlight consists of various amounts of the entire spectrum of visible light, in addition to ultraviolet and infrared radiation. Because sunlight contains more red than far-red light, however, when a plant is exposed to sunlight, the level of Pfr increases. During the night, the level of Pfr slowly decreases as Pfr, which is less stable than Pr, is degraded. The importance of phytochrome to plants cannot be overemphasized. Timing of day length and darkness is the most reliable way for plants to measure the change from one season to the next. This measurement, which synchronizes the stages of plant development, is crucial for survival, particularly in environments where the climate has an annual pattern of favorable and unfavorable seasons.

ing the equilibrium between the Pr and Pfr forms, particularly of phyB. This signal triggers a series of responses that cause the shade-avoiding plant, which is adapted to full-light environments, to grow taller or flower earlier. FIGURE 38-19 shows an interesting application of additional (reflected) far-red light without the reduction in incoming sunlight that occurs in shade avoidance.

Plants sense the proximity of nearby plants, which are potential competitors, and react by changing the way they grow and develop. Many plants, from small herbs to large trees, compete for light, a response known as shade avoidance, in which plants tend to grow taller when closely surrounded by other plants. If successful, the shade-avoiding plant projects its new growth into direct sunlight and thereby increases its chances of survival. Since the 1970s, botanists have recognized the environmental factor that triggers shade avoidance: plants perceive changes in the ratio of red to far-red light that result from the presence of nearby plants. The leaves of neighboring plants absorb much more red light than far-red light. (The green pigment chlorophyll strongly absorbs red light during photosynthesis.) In a densely plant-populated area, the ratio of red light to far-red light (r/fr) decreases, affect-

Cary Wolinsky

Competition for sunlight among shade-avoiding plants involves phytochrome

FIGURE 38-19 Colored mulches and plant growth processes The red mulch reflects far-red light from the ground to the tomato plants. The plants, absorbing the extra far-red light, react as if nearby plants are present. Their aboveground growth is greater, and their fruits ripen earlier. Manipulating mulch colors also modifies yield, flavor, and nutrient content.

When a plant is using many of its resources for stem elongation, it has fewer resources for new leaves and branches, storage tissues, and reproductive tissues. However, for a shade-avoiding plant that is shaded by its neighbors, a rapid increase in stem length is advantageous, because once this plant is taller than its neighbors, it obtains a larger share of unfiltered sunlight.

Phytochrome is involved in other responses to light, including germination Phytochrome is involved in the light requirement that some seeds have for germination. Seeds with a light requirement must be exposed to light containing red wavelengths. Exposure to red light converts Pr to Pfr, and germination occurs. Many temperate forest species with small seeds require light for germination. (Larger seeds generally do not have a light requirement.) This adaptation enables the seeds to germinate at the optimal time. During early spring, sunlight, including red light, penetrates the bare branches of overlying deciduous trees and reaches the soil between the trees. As spring temperatures warm, the seeds on the soil absorb red light and germinate. During their early growth, the newly germinated seedlings do not have to compete with the taller trees for sunlight. Other physiological functions under the influence of phytochrome include sleep movements in leaves (discussed later); shoot dormancy; leaf abscission; and pigment formation in flowers, fruits, and leaves.

Phytochrome acts by signal transduction Some phytochrome-induced responses are very rapid and shortterm, whereas others are slower and long-term. The rapid responses probably involve reversible changes in the properties of membranes, altering cell ionic balances. Red light, for example, causes potassium ion (K+) channels to open in cell membranes involved in plant movements caused by changes in turgor. Exposure to far-red light causes the K+ channels to close. Slower phytochrome-induced responses involve the selective regulation of transcription of numerous genes. For example, phytochrome activates transcription of the gene for the small subunit of rubisco, an enzyme involved in photosynthesis (see Chapter 9). Each phytochrome molecule consists of a protein attached to a light-absorbing photoreceptor. Biologists hypothesize that the absorption of light by the photoreceptor portion of phytochrome elicits a change in the shape of the protein portion of the phytochrome molecule. This change, in turn, triggers one or more signal transduction pathways. One important research tool in studying phytochrome signal transduction is mutant plants that respond as if they were exposed to a particular light stimulus even if they were not. Research indicates that the active form of phytochrome moves from the cytoplasm into the nucleus, where it affects gene expression by activating a transcription factor. When activated, the transcription factor, which binds to the promoter of a gene, either turns on or represses transcription that leads to protein synthesis. In one such pathway, inactive phytochrome (Pr) in the cytoplasm absorbs red light and is converted into the active form, Pfr, which moves into the nucleus. There, phytochrome binds to PIF3 (phytochrome-interacting factor 3), a transcription factor that is

already bound to the promoters of light-responsive genes, thereby activating (or repressing) their transcription. The signal transduction pathway is shut down by far-red light, which is absorbed by the Pfr in the Pfr–PIF3 complex in the nucleus. When Pfr is converted to Pr, it dissociates from PIF3. The signal transduction pathway just described is not the end of the story. Phytochrome does not regulate just one but numerous light-responsive signaling pathways that affect gene expression. Also, we do not understand the specifics of how the transition from Pr to Pfr participates in so many signaling cascades.

Light influences circadian rhythms Almost all organisms, including plants, animals, fungi, eukaryotic microorganisms, and many bacteria, appear to have an internal timer, or biological clock, that approximates a 24-hour cycle, the time it takes for Earth to rotate around its own axis. These internal molecular cycles are known as circadian rhythms (from the Latin circum, “around,” and diurn, “daily”). Circadian rhythms help an organism detect the time of day, whereas photoperiodism enables a plant to detect the time of year. Why do plants and other organisms exhibit daily circadian rhythms? Predictable environmental changes, such as sunrise and sunset, occur during the course of each 24-hour period. These predictable changes may be important to an individual organism, causing it to change its physiological activities or its behavior (in the case of animals). Researchers think that circadian rhythms help an organism synchronize repeated daily activities so that they occur at the appropriate time each day. If, for example, an insectpollinated flower does not open at the time of day that pollinating insects are foraging for food, reproduction will be unsuccessful. Circadian rhythms in plants affect such biological events as gene expression, the timing of photoperiodism (that is, of seasonal reproduction), the rate of photosynthesis, and the opening and closing of stomata. Sleep movements observed in the common bean and other plants are another example of a circadian rhythm (FIG. 38-20). During the day, bean leaves are horizontal, possibly for optimal light absorption, but at night the leaves fold down or up, a movement that orients them perpendicular to their daytime position. The biological significance of sleep movements is unknown at this time. When constant environmental conditions are maintained, circadian rhythms such as sleep movements repeat every 20 to 30 hours, at least for several days. In nature, the rising and setting of the sun reset the biological clock so that the cycle repeats every 24 hours. What happens if a plant’s circadian clock is mutated so that it is not resynchronized by the day–night cycle? In 2005, biologists at the University of Cambridge in England identified mutants of the model plant Arabidopsis in which the circadian clock is not synchronized to match the external day–night cycle. The mutant plants contained less chlorophyll, fixed less carbon by photosynthesis, grew more slowly, and had less of a competitive advantage than Arabidopsis plants with a normal circadian clock. For Arabidopsis and many other plants, two photoreceptors— the red light–absorbing phytochrome and the blue/ultraviolet-A light–absorbing cryptochrome—are implicated in resetting the biological clock. Certain amino acid sequences of the protein portion of phytochrome are homologous with amino acid sequences

Cengage

Cengage

(a) Leaf position at noon in a bean (Phaseolus vulgaris) seedling.

The plant also sends signals from the site of localized infection, leading to systemic acquired resistance (SAR), a buildup of long-lasting, enhanced resistance throughout the plant. The signal travels in the phloem. Systemic acquired resistance is broad spectrum—that is, SAR is a heightened defense against many types of pathogens, not just the one that caused the initial infection. Plant biologists continue to discover new hormonelike signaling molecules. Many of these signaling molecules are involved in the defensive responses of plants to disease organisms and insects. Here we briefly consider two groups—jasmonic acid and methyl salicylate.

(b) Leaf position at midnight.

FIGURE 38-20 Animated Sleep movements

of clock proteins in fruit flies, fungi, mammals, and bacteria; this molecular evidence strongly supports the circadian-clock role of phytochrome. The evidence for cryptochrome as a clock protein is also convincing. First discovered in plants, cryptochrome counterparts are found in the fruit fly and mouse biological-clock proteins. Possibly both photoreceptors are involved in resetting the biological clock in plants; researchers have evidence that phytochrome and cryptochrome sometimes interact to regulate similar responses.

Jasmonic acid activates several plant defenses Jasmonic acid affects several plant processes, such as pollen development, root growth, fruit ripening, and senescence. This lipidderived plant hormone, which is structurally similar to prostaglandins in animals, is also produced in response to the presence of insect pests and disease-causing organisms. Jasmonic acid triggers the production of enzymes that confer an increased resistance against herbivorous (plant-eating) insects. For example, caterpillar-infested tomato plants release volatile jasmonic acid into the air that attracts natural caterpillar enemies, such as parasitic wasps; the wasps lay their eggs on the caterpillars’

Review ■ ■ ■

What is phytochrome? What are two roles of phytochrome? What are the steps in phytochrome signal transduction? In what process is cryptochrome involved?

38.4 RESPONSES TO HERBIVORES AND PATHOGENS ■ ■ LEARNING OBJECTIVE 9 Distinguish between the hypersensitive response and systemic acquired

Plants lack an immune system such as is found in humans and other mammals, but plants can defend themselves from attack by pathogenic and herbivorous enemies. Each plant cell has an innate immune system to fight against local infections. When a molecule produced by a pathogen or herbivore binds to a receptor in a plant cell, it triggers a signal transduction pathway. As a result, a hypersensitive response occurs that seals off the infected area, causing it to die. These necrotic lesions, or dead areas, prevent or slow the spread of the initial infection (FIG. 38-21).

© Nigel Cattlin/Alamy

resistance.

FIGURE 38-21 Hypersensitive response The necrotic lesions on these oak leaves were produced in response to invasion by a fungal pathogen.

bodies. When the eggs hatch, the wasp larvae consume, and ultimately kill, the host insects. In one study, treating plants with jasmonic acid increased parasitism on caterpillar pests twofold over control fields in which the plants did not have jasmonic acid applications. Jasmonic acid may be of practical value in controlling certain insect pests without the use of chemical pesticides. Biologists have made progress in identifying key aspects of the jasmonic acid signaling pathway. Interestingly, the jasmonic acid signaling pathway includes ubiquitin, which was discussed earlier in the chapter in relation to auxin (see Fig. 38-4). As in the auxin signaling pathway, the ubiquitin tags in the jasmonic acid signaling pathway cause degradation of transcription repressors, thereby altering gene expression.

Methyl salicylate may induce systemic acquired resistance For centuries people chewed willow (Salix) bark to treat headaches and other types of pain. Salicylic acid was first extracted from willow bark and is chemically related to aspirin (acetylsali-

■

cylic acid). More recently, biologists have shown that salicylic acid helps plants defend against insect pests and pathogens such as viruses. When a plant is under attack, the concentration of salicylic acid increases, and it spreads systemically throughout the plant. Based on current research, biologists think that methyl salicylate, a derivative of salicylic acid also known as oil of wintergreen, binds to a cell receptor, triggering a signal transduction pathway that switches on genes. These genes code for proteins that establish systemic acquired resistance to fight infection and promote wound healing. Plants also use methyl salicylate, which is volatile, to signal nearby plants. Tobacco plants infected with tobacco mosaic virus release methyl salicylate into the air. When nearby healthy plants receive the airborne chemical signal, they begin synthesizing antiviral proteins that enhance their resistance to the virus.

Review ■ ■

What is the hypersensitive response? What is systemic acquired resistance?

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

38.1 (page 804)

fused from the coleoptile tip into the agar and, later, from the agar into the decapitated coleoptile. Went named this substance auxin.

1 Describe phototropism, gravitropism, and thigmotropism. ■

Tropisms are directional growth responses. Phototropism is growth in response to the direction of light. Gravitropism is growth in response to the influence of gravity. Thigmotropism is growth in response to contact with a solid object. Learn more about phototropism and gravitropism by clicking on the figures in CengageNOW.

38.2 (page 805) 2 Describe a general mechanism of action for plant hormones, using auxin as your example. Plants produce and respond to hormones, organic molecules that act as highly specific chemical signals to elicit a variety of responses that regulate growth and development. Many plant hormones bind to enzyme-linked receptors located in ■ the plasma membrane; the hormone binds to the receptor, triggering an enzymatic reaction of some sort. The receptor for auxin, which is located in the cytosol or nucleus, binds to auxin. This catalyzes the attachment of ubiquitin to a repressor protein, targeting it for destruction. When genes repressed by those proteins are then expressed, changes in cell growth and development result. 3 Describe early auxin experiments involving phototropism. ■ Darwin and his son performed phototropism experiments in the 1870s on grass seedlings. When they covered the coleoptile tip, the plant did not bend. When they removed the coleoptile tip, bending did not occur. When the bottom of the coleoptile was shielded from the light, the coleoptile bent. The Darwins concluded that some substance is transmitted from the upper to the lower part that causes the plant to bend. In the 1920s, Frits Went isolated the phototropic hormone from oat ■ coleoptiles by removing the coleoptile tips and placing them on agar blocks. Normal growth resumed when he put one of these agar blocks squarely on a decapitated coleoptile; the substance had dif■

See the auxin experiments in action by clicking on the figure in CengageNOW.

4 List several ways that each of these hormones affects plant growth and development: auxins, gibberellins, cytokinins, ethylene, abscisic acid, and brassinosteroids. ■ Auxin is involved in cell elongation; tropisms; apical dominance, the inhibition of axillary buds by the apical meristem; and fruit development. Auxin also stimulates root development on stem cuttings. ■ Gibberellins are involved in stem elongation, flowering, and germination. Cytokinins promote cell division and differentiation; delay se■ nescence, the natural aging process; and interact with auxin and ethylene in apical dominance. Cytokinins induce cell division in tissue culture, a technique in which cells are isolated from plants and grown in a nutrient medium. ■ Ethylene plays a role in ripening fruits; apical dominance; leaf abscission; wound response; thigmomorphogenesis, a developmental response to mechanical stressors such as wind; and senescence.

■

Abscisic acid is an environmental stress hormone involved in stomatal closure caused by water stress and in seed dormancy, a temporary state of reduced physiological activity.

■

Plant steroids known as brassinosteroids are involved in several aspects of plant growth and development, such as cell division, cell elongation, light-induced differentiation, seed germination, and vascular development.

cleus. There, phytochrome binds to the transcription factor PIF3 (phytochrome-interacting factor) and activates or represses the transcription of light-responsive genes. Learn more about phytochrome by clicking on the figure in CengageNOW.

38.3 (page 813) 5 Explain how varying amounts of light and darkness induce flowering. ■

Photoperiodism is any response of plants to the duration and timing of light and dark. Flowering is a photoperiodic response in many plants. Short-day plants detect the lengthening nights of late summer or fall and flower at that time. Long-day plants detect the shortening nights of spring and early summer and flower at that time. Intermediate-day plants flower when exposed to days and nights of intermediate length. Learn more about photoperiodism by clicking on the figure in CengageNOW.

6 Describe the role of phytochrome in flowering, including a brief discussion of phytochrome signal transduction. ■ The photoreceptor in photoperiodism is phytochrome, a family of about five blue-green pigments. Each type of phytochrome has two forms, Pr and Pfr, named by the wavelength of light they absorb. Pfr is the active form, triggering or inhibiting physiological responses such as flowering, shade avoidance, and a light requirement for germination. ■ The pathway in phytochrome signal transduction begins when inactive phytochrome in the cytoplasm absorbs red light and is converted into the active form, Pfr, which moves into the nu-

7 Define circadian rhythm and give an example. A circadian rhythm is a regular period in an organism’s growth or activities that approximates the 24-hour day and is reset by the rising and setting of the sun. Two examples of circadian rhythms are the opening and closing of stomata and sleep movements. 8 Distinguish between phytochrome and cryptochrome. ■ Both phytochrome and cryptochrome are photoreceptors that sometimes interact to regulate similar responses, such as resetting the biological clock. Phytochrome strongly absorbs red light, whereas cryptochrome absorbs blue and ultraviolet-A light. ■

38.4 (page 817) 9 Distinguish between the hypersensitive response and systemic acquired resistance. ■ Each plant cell has an innate immune system to fight against local infections. A hypersensitive response seals off the infected area, causing it to die. These dead areas prevent or slow the spread of the initial infection. ■ Systemic acquired resistance (SAR) is a buildup of long-lasting, enhanced resistance throughout the plant. Systemic acquired resistance is broad spectrum (against many types of pathogens).

T E S T YO U R U N D E R S TA N D I N G 1. In the signal transduction process for the hormone auxin, the molecule ubiquitin (a) absorbs blue light (b) becomes phosphorylated (c) tags certain proteins for destruction (d) interacts antagonistically with gibberellins (e) binds to a receptor in the plant cell’s plasma membrane 2. When you prune shrubs to make them “bushier”—that is, to prevent apical dominance—you are affecting the distribution and action of which plant hormone? (a) auxin (b) jasmonic acid (c) salicylic acid (d) ethylene (e) abscisic acid 3. Research on a fungal disease of rice provided the first clues about the plant hormone (a) auxin (b) gibberellin (c) cytokinin (d) ethylene (e) abscisic acid 4. This plant hormone interacts with auxin during the formation of plant organs in tissue culture. (a) florigen (b) gibberellin (c) cytokinin (d) ethylene (e) abscisic acid 5. The stress hormone that helps plants respond to drought is (a) auxin (b) gibberellin (c) cytokinin (d) ethylene (e) abscisic acid 6. This hormone promotes seed dormancy. (a) auxin (b) gibberellin (c) cytokinin (d) ethylene (e) abscisic acid 7. The three classes of photoreceptors that enable plants to sense the presence and duration of light are (a) phytochrome, cryptochrome, and circadian rhythms (b) chlorophyll, photosynthesis, and photoperiodism (c) phytochrome, photoperiodism, and chlorophyll (d) phytochrome, cryptochrome, and phototropin (e) phototropin, photosynthesis, and photoperiodism

8. A plant’s response to the relative amounts of daylight and darkness is (a) apical dominance (b) bolting (c) gravitropism (d) photoperiodism (e) phototropism 9. Pfr, the active state of _____, forms when red light is absorbed. (a) photosystem I (b) phytochrome (c) far-red light (d) phototropin (e) cryptochrome 10. Which of the following represents the correct order in the phytochrome signal transduction pathway? 1. red light 2. light-responsive gene is switched on (or off ) 3. movement of Pfr to nucleus 4. conversion of Pr to Pfr 5. formation of PFr–PIF3 complex that is bound to promoter region (a) 1, 3, 5, 4, 2 (b) 1, 5, 3, 2, 4 (c) 1, 2, 3, 4, 5 (d) 1, 4, 3, 2, 5 (e) 1, 4, 3, 5, 2 11. Which photoreceptor(s) is/are implicated in resetting the biological clock? (a) phytochrome only (b) cryptochrome only (c) phototropin only (d) cryptochrome and gibberellin (e) phytochrome and cryptochrome 12. Which signaling molecule triggers the release of volatile substances that attract parasitic wasps to plant-eating caterpillars? (a) phytochrome (b) jasmonic acid (c) auxin (d) methyl salicylate (e) gibberellin

CRITICAL THINKING

2. If you transplanted the short-day plant discussed in question 1 to the tropics, would it flower? Explain your answer. 3. EVOLUTION LINK. What adaptive advantages are conferred on a plant whose stems are positively phototropic and whose roots are positively gravitropic? 4. EVOLUTION LINK. Explain why a plant with a mutation in one of its phytochrome genes that affects flowering time would be at an evolutionary disadvantage. 5. ANALYZING DATA. Examine the figure (right). When the day length is 10 hours and the night length is 14 hours, what percent of the plants flower? When the day length is 16 hours and the night

length is 8 hours, what percent of the plants flower? What is the critical night length for this plant?

Percent flowering

1. Predict whether flowering in a short-day plant with a minimum critical night length of 14 hours would be expected to occur in the following situations. Explain each answer. (a) The plant is exposed to 15 hours of daylight and 9 hours of uninterrupted darkness. (b)The plant is exposed to 9 hours of daylight and 15 hours of darkness. (c) The plant is exposed to 9 hours of daylight and 15 hours of darkness, with a 10-minute exposure to red light in the middle of the night.

100

0 Day length

10

12

14

16

Night length

14

12

10

8

6. SCIENCE, TECHNOLOGY, AND SOCIETY. Do you think our expanding knowledge of the molecular basis of plant signaling molecules could improve agricultural productivity in the future? Explain your answer. Additional questions are available in CengageNOW at www.cengage.com/ login.

39

Animal Structure and Function: An Introduction

Fritz Polking/Dembinsky Photo Associates

Larger body size does not mean bigger cells. The cells of the Yacare caiman (Caiman yacare) and the flambeau butterfly (Dryas julia) on its head are all about the same size. The caiman is larger because its genes specify that its body consists of a larger number of cells.

KEY CONCEPTS 39.1 Cells make up tissues, various types of tissues make up organs, and tissues and organs working together make up organ systems. The main types of tissues found in animals are epithelial, connective, muscle, and nervous tissues. 39.2 Homeostatic mechanisms are mainly negative feedback systems that maintain a relatively stable internal environment.

39.3 Thermoregulation is the process of maintaining homeostasis of body temperature, despite changes in surrounding (or internal) temperature.

A

nimal groups are dramatically diverse, with radically different body structures. For example, consider how different the caiman and

the butterfly are, not only in size but also in body form and lifestyle (see photograph). Despite their differences, animal groups share many characteristics, including their relatively large size. Why are most animals larger than bacteria, protists, and fungi? The answer may be related to ecological niches, which are the functional roles of a species within a community. By the time animals evolved, bacteria, protists, and fungi already occupied most available ecological niches. For new species to succeed, they had to displace others from a niche or adapt to a new one. Success in a new niche required a new body plan, and new body plans often required and accommodated larger size. Increased size also provided more opportunity for capturing food. Predators are typically larger than their prey. To grow larger than their bacterial and protist competitors, animals had to be multicellular. Recall that the size of a single cell is limited by the ratio of its surface area (plasma membrane) to its volume (see Chapter 4). The plasma membrane needs to be large enough relative to the cell’s volume to permit passage of materials into and out of the cell so that the

conditions necessary for life can be maintained. In a multicellular animal, each cell has a large enough surface-area-to-volume ratio to effectively regulate its internal environment. Individual cells live and die and are replaced while the organism continues to maintain itself and thrive. The number of cells, not their individual sizes, is mainly responsible for the size of an animal. In unicellular organisms, such as bacteria and many protists, the single cell carries on all the activities necessary for life. Recall that unicellular and small, flat organisms depend on diffusion for many life processes, including gas exchange and disposal of metabolic wastes. One reason they can be small is that they do not require complex organ systems. In this chapter we focus on the basic form and function of the animal body. Anatomy is the study of an organism’s structure. Physiology is the study of how the body functions. Structure and function are closely linked at every level of organization. We describe the types and functions of tissues and the principal organ systems of animals. We also discuss the important concept of homeostasis, using regulation of body temperature as an example. In the following chapters, we discuss how organ systems work together to maintain homeostasis, as the animal carries out its many life processes.

39.1 TISSUES, ORGANS, AND ORGAN SYSTEMS ■ ■ LEARNING OBJECTIVES 1 2 3 4 5 6

Compare the structure and function of the four main kinds of animal tissues: epithelial, connective, muscle, and nervous tissues. Compare the main types of epithelial tissue and describe their functions. Compare the main types of connective tissue and describe their functions. Contrast the three types of muscle tissue and describe their functions. Relate the structure of the neuron to its function. Briefly describe the organ systems of a mammal and summarize the functions of each organ system.

In a multicellular organism, cells specialize to perform specific tasks. Recall from Chapter 1 that cells organize to form tissues, and tissues associate to form organs such as the heart or stomach. Groups of tissues and organs make up the organ systems of a complex organism. Billions of cells may be organized to form the tissues, organs, and organ systems of a complex animal. A tissue consists of a group of closely associated, similar cells that carry out specific functions. Biologists classify animal tissues as epithelial, connective, muscle, or nervous tissue. Classification of tissues depends on their structure and origin. Each kind of tissue is composed of cells with characteristic sizes, shapes, and arrangements; and each type of tissue is specialized to perform a specific function or group of functions. For example, some tissues are specialized to transport materials, whereas others contract, enabling the animal to move. Still others secrete hormones that regulate

metabolic processes. Structure and function are closely linked at every level of organization. As we discuss each tissue type, notice the relationship between its form and its function.

Epithelial tissues cover the body and line its cavities Epithelial tissue (also called epithelium) consists of cells fitted tightly together to form a continuous layer, or sheet, of cells. One surface of the sheet is typically exposed because it covers the body (outer layer of the skin) or lines a cavity, such as the lumen (the cavity in a hollow organ) of the intestine. The other surface of an epithelial layer attaches to the underlying tissue by a noncellular basement membrane consisting of tiny fibers and nonliving polysaccharide material that the epithelial cells produce. Epithelial tissue forms the outer layer of the skin and the linings of the digestive, respiratory, excretory, and reproductive tracts. As a result, everything that enters or leaves the body must cross at least one layer of epithelium. Food taken into the mouth and swallowed is not really “inside” the body until it is absorbed through the epithelium of the gut and enters the blood. To a large extent, the permeabilities of the various epithelial tissues regulate the exchange of substances between the different parts of the body, as well as between the animal and the external environment. Epithelial tissues perform many functions, including protection, absorption, secretion, and sensation. The epithelial layer of the skin, the epidermis, covers the entire body and protects it from mechanical injury, chemicals, bacteria, and fluid loss. The epithelial tissue lining the digestive tract absorbs nutrients and water into the body. Some epithelial cells form glands that secrete cell products such as hormones, enzymes, or sweat. Other epithelial cells are sensory receptors that receive information from the environment. For example, epithelial cells in taste buds and in the nose specialize as chemical receptors. TABLE 39-1 illustrates the main types of epithelial tissue, indicates their locations in the body, and describes their functions (pages 824–825). We can distinguish three types of epithelial cells on the basis of shape. Squamous epithelial cells are thin, flat cells shaped like flagstones. Cuboidal epithelial cells are short cylinders that from the side appear cube-shaped, like dice. Actually, each cuboidal cell is typically hexagonal in cross section, making it an eight-sided polyhedron. Columnar epithelial cells look like columns or cylinders when viewed from the side. The nucleus is usually located near the base of the cell. Viewed from above or in cross section, these cells often appear hexagonal. On its free surface, a columnar epithelial cell may have cilia that beat in a coordinated way, moving materials over the tissue surface. Most of the upper respiratory tract is lined with ciliated columnar epithelium that moves particles of dust and other foreign material away from the lungs. Epithelial tissue is also classified by number of layers. Simple epithelium is composed of one layer of cells. It is usually located where substances are secreted, excreted, or absorbed, or where materials diffuse between compartments. For example, simple squamous epithelium lines the air sacs in the lungs. The structure of this thin tissue allows diffusion of gases in and out of air sacs. Stratified epithelium, which has two or more layers, protects underlying tissues. For example, stratified squamous epithelium,

which makes up the outer layer of your skin, is continuously sloughed off during normal wear and tear. It must also continuously regenerate. The cells of pseudostratified epithelium falsely appear layered. Although all its cells rest on a basement membrane, not every cell extends to the exposed surface of the tissue. This arrangement gives the impression of two or more cell layers. Some of the respiratory passageways are lined with pseudostratified epithelium equipped with cilia. The lining of blood and lymph vessels is called endothelium. Endothelial cells have a different embryonic origin from “true” epithelium. However, these cells are structurally similar to squamous epithelial cells and can be included in that category. Glands are made of epithelial cells A gland consists of one or more epithelial cells specialized to produce and secrete a product such as sweat, milk, mucus, wax, saliva, hormones, or enzymes (FIG. 39-1). Epithelial tissue lining the cavities and passageways of the body typically has some specialized mucus-secreting cells called goblet cells. The mucus lubricates these surfaces, offers protection, and facilitates the movement of materials. Glands are classified as exocrine or endocrine. Exocrine glands, like goblet cells and sweat glands, secrete their products onto a free epithelial surface, typically through a duct (tube). Endocrine glands lack ducts. These glands release their products, called hormones, into the interstitial fluid (tissue fluid) or blood; hormones are typically transported by the cardiovascular system. (Endocrine glands are discussed in Chapter 50.)

Unicellular glands (goblet cells)

Cilia

Basement membrane

(a) Goblet cells. These unicellular glands secrete mucus.

Skin

(b) Sweat gland. These simple glands consist of coiled tubes. Their walls are constructed of simple cuboidal epithelium.

(c) Parotid salivary gland. Compound glands, like the parotid, have branched ducts.

FIGURE 39-1 Glands

Epithelial cells form membranes A gland consists of one or more epithelial cells. An epithelial membrane consists of a sheet of epithelial tissue and a layer of underlying connective tissue. Types function of each kind of connective tissue are determined in part of epithelial membranes include mucous membranes and serous by the structure and properties of the intercellular substance. membranes. A mucous membrane, or mucosa, lines a body cavConnective tissue typically contains three types of fibers: collaity that opens to the outside of the body, such as the digestive or gen, elastic, and reticular. Collagen fibers, the most numerous type, respiratory tract. Goblet cells in the epithelial layer secrete mucus are made of collagens, a group of fibrous proteins found in all animals that lubricates the tissue and protects it from drying. (see Fig. 3-22b). Collagens are the most abundant proteins in mamA serous membrane lines a body cavity that does not open mals, accounting for about 25% of their total protein mass. Collagen is to the outside of the body. It consists of simple squamous epivery tough (meat is tough because of its collagen content). The tensile thelium over a thin layer of loose connective tissue. This type of strength (ability to stretch without tearing) of collagen fibers is commembrane secretes fluid into the cavity it lines. Examples of serous parable to that of steel. Collagen fibers are wavy and flexible, allowing membranes are the pleural membranes lining the pleural cavities them to remain intact when tissue is stretched. around the lungs and the pericardial membranes lining the pericarElastic fibers branch and fuse to form networks. They can be dial cavity around the heart. stretched by a force and then (like a stretched rubber band) return to their original size and shape when the force is removed. Elastic Connective tissues support fibers, composed of the protein elastin, are an important compoother body structures nent of structures that must stretch. Almost every organ in the body has a framework of connective Reticular fibers are very thin, branched fibers that form delitissue that supports and cushions it. Compared to epithelial tiscate networks joining connective tissues to neighboring tissues. sues, connective tissues contain relatively few cells. Its cells are Reticular fibers consist of collagen and some glycoprotein. embedded in an extensive intercellular substance consisting of The cells of various kinds of connective tissues differ in their threadlike, microscopic fibers scattered throughout a matrix, a shapes and structures and in the kinds of fibers and matrices they thin gel of polysaccharides that the cells secrete. The nature and secrete. Fibroblasts are connective tissue cells that produce the

TABLE 39-1

Ephithelial Tissues Nuclei of squamous epithelial cells

Ed Reschke

Simple Squamous Epithelium Main Locations Air sacs of lungs; lining of blood vessels Functions Passage of materials where little or no protection is needed and where diffusion is major form of transport Description and Comments Cells are flat and arranged as single layer

25 μm LM of simple squamous epithelium.

Nuclei of cuboidal epithelial cells

Lumen of tubule

Ed Reschke

Simple Cuboidal Epithelium Main Locations Linings of kidney tubules; gland ducts Functions Secretion and absorption Description and Comments Single layer of cells; LM shows cross section through tubules; from the side each cell looks like a short cylinder; some have microvilli for absorption

25 μm LM of simple cuboidal epithelium.

Goblet cell

Nuclei of columnar cells

Ed Reschke

Simple Columnar Epithelium Main Locations Linings of much of digestive tract and upper part of respiratory tract Functions Secretion, especially of mucus; absorption; protection; moves layer of mucus Description and Comments Single layer of columnar cells; sometimes with enclosed secretory vesicles (in goblet cells); highly developed Golgi complex; often ciliated

25 μm LM of simple columnar epithelium.

(continued)

TABLE 39-1

Ephithelial Tissues (continued)

Ed Reschke

Stratified Squamous Epithelium Main Locations Skin; mouth lining; vaginal lining Functions Protection only; little or no absorption or transit of materials; outer layer continuously sloughed off and replaced from below Description and Comments Several layers of cells, with only the lower ones columnar and metabolically active; division of lower cells causes older ones to be pushed upward toward surface, becoming flatter as they move

50 μm LM of stratified squamous epithelium.

Cilia

Epithelial cells

Pseudostratified Epithelium Main Locations Some respiratory passages; ducts of many glands Functions Secretion; protection; moves layer of mucus Description and Comments Ciliated, mucus-secreting, or with microvilli; comparable in many ways to columnar epithelium except that not all cells are the same height; so, though all cells contact the same basement membrane, the tissue appears stratified

Nuclei

Ed Reschke

Basement membrane

25 μm LM of pseudostratified columnar epithelium, ciliated.

fibers, as well as the protein and carbohydrate complexes, of the matrix. Fibroblasts release protein components that become arranged to form the characteristic fibers. These cells are especially active in developing tissues and are important in healing wounds. As tissues mature, the number of fibroblasts decreases and they become less active. Macrophages, the body’s scavenger cells, commonly wander through connective tissues, cleaning up cell debris and phagocytosing foreign matter, including bacteria.

Some of the main types of connective tissue are (1) loose and dense connective tissues; (2) elastic connective tissue; (3) reticular connective tissue; (4) adipose tissue; (5) cartilage; (6) bone; and (7) blood, lymph, and tissues that produce blood cells. These tissues vary widely in their structural details and in the functions they perform (TABLE 39-2). Loose connective tissue is the most widely distributed connective tissue in the vertebrate body. Together with adipose tissue,

TABLE 39-2 Collagen fibers

Connective Tissues Nuclei of fibroblasts

Ed Reschke

Elastic fibers

Loose Connective Tissue Main Locations Everywhere that support must be combined with elasticity, such as subcutaneous tissue (the layer of tissue beneath the dermis of the skin) Functions Support; reservoir for fluid and salts Description and Comments Fibers produced by fibroblast cells embedded in semifluid matrix and mixed with miscellaneous other cells

50 μm LM of loose connective tissue.

Nucleus of fibroblast

Ed Reschke

Collagen fibers

Dense Connective Tissue Main Locations Tendons; many ligaments; dermis of skin Functions Support; transmits mechanical forces Description and Comments Collagen fibers may be regularly or irregularly arranged

25 μm LM of dense connective tissue.

Elastic fibers

Ed Reschke

Elastic Connective Tissue Main Locations Structures that must both expand and return to their original size, such as lung tissue and large arteries Function Confers elasticity Description and Comments Branching elastic fibers interspersed with fibroblasts

50 μm LM of elastic connective tissue.

Reticular fibers

Ed Reschke

Reticular Connective Tissue Main Locations Framework of liver; lymph nodes; spleen Function Support Description and Comments Consists of interlacing reticular fibers

50 μm LM of reticular connective tissue.

(continued)

TABLE 39-2

Connective Tissues (continued)

Cengage

Adipose Tissue Main Locations Subcutaneous layer; pads around certain internal organs Functions Food storage; insulation; supports organs such as mammary glands, kidneys Description and Comments Fat cells are star shaped at first; fat droplets accumulate until typical ringshaped cells are produced 50 μm LM of adipose tissue.

Chondrocytes

Lacuna

Intercellular substance

Ed Reschke

Cartilage Main Locations Supporting skeletons in sharks and rays; ends of bones in mammals and some other vertebrates; supporting rings in walls of some respiratory tubes; tip of nose; external ear Function Flexible support Description and Comments Cells (chondrocytes) separated from one another by intercellular substance; cells occupy lacunae 50 μm LM of cartilage. Lacunae

Haversian canal

Matrix

Cengage

Bone Main Locations Forms skeletal structure in most vertebrates Functions Supports and protects internal organs; calcium reservoir; skeletal muscles attach to bones Description and Comments Osteocytes in lacunae; in compact bone, lacunae embedded in lamellae, concentric circles of matrix surrounding Haversian canals 50 μm LM of bone. Red blood cells

White blood cells

Ed Reschke

Blood Main Locations Within heart and blood vessels of circulatory system Functions Transports oxygen, nutrients, wastes, and other materials Description and Comments Consists of cells dispersed in fluid intercellular substance (plasma)

25 μm LM of blood.

loose connective tissue forms the subcutaneous (below the skin) layer that attaches skin to the muscles and other structures beneath. Nerves, blood vessels, and muscles are wrapped in loose connective tissue. This tissue also forms a thin filling between body parts and serves as a reservoir for fluid and salts. Loose connective tissue consists of fibers running in all directions through a semifluid matrix. Its flexibility permits the parts it connects to move. Dense connective tissue, found in the dermis (lower layer) of the skin, is very strong, but is somewhat less flexible than loose connective tissue. Collagen fibers predominate. Tendons, the cords that connect muscles to bones, and ligaments, the cables that connect bones to one another, consist of dense connective tissue in which collagen bundles are arranged in a definite pattern. Elastic connective tissue consists mainly of bundles of parallel elastic fibers. This tissue is found in structures that must expand and then return to their original size, such as lung tissue and the walls of large arteries. Reticular connective tissue is composed mainly of interlacing reticular fibers. It forms a supporting internal framework in many organs, including the liver, spleen, and lymph nodes. The cells of adipose tissue store fat and release it when fuel is needed for cellular respiration. Adipose tissue is found in the subcutaneous layer and in tissue that cushions internal organs. The supporting skeleton of a vertebrate is made of cartilage, or of both cartilage and bone. Cartilage is the supporting skeleton in the embryonic stages of all vertebrates. In most vertebrates, bone replaces cartilage during development. However, cartilage remains in some supporting structures. For example, in humans, cartilage is found in the external ear, the supporting rings in the walls of the respiratory passageways, the tip of the nose, the ends of some bones, and the discs that serve as cushions between the vertebrae. Cartilage is firm yet elastic. Its cells, called chondrocytes, secrete a hard, rubbery matrix that surrounds them. They also secrete collagen fibers, which become embedded in the matrix and strengthen it. Chondrocytes eventually come to lie, singly or in groups of two or four, in small cavities in the matrix called lacunae. These cells remain alive and are nourished by nutrients and oxygen that diffuse through the matrix. Cartilage tissue lacks nerves, lymph vessels, and blood vessels. Bone, the main vertebrate skeletal tissue, is like cartilage in that it consists mostly of matrix material. The bone cells, called osteocytes, are contained within lacunae. Osteocytes secrete and maintain the matrix (FIG. 39-2). Unlike cartilage, however, bone is a highly vascular tissue, with a substantial blood supply. Osteocytes communicate with one another and with capillaries by tiny channels (canaliculi) that contain long cytoplasmic extensions of the osteocytes. A typical bone has an outer layer of compact bone surrounding a filling of spongy bone. Compact bone consists of spindle-shaped units called osteons. Within each osteon, osteocytes are arranged in concentric layers of matrix called lamellae. In turn, the lamellae surround central microscopic channels known as Haversian canals, through which capillaries and nerves pass. Bones are amazingly light and strong. Calcium salts of bone render the matrix very hard, and collagen prevents the bony matrix from being overly brittle. Most bones have a large, central marrow cavity that contains a spongy tissue called marrow. Yellow mar-

row consists mainly of fat. Red marrow is the connective tissue in which blood cells are produced. We discuss bone in more detail in Chapter 40. Blood and lymph are circulating tissues that help other parts of the body communicate and interact. Like other connective tissues, they consist of specialized cells dispersed in an intercellular substance. In mammals, blood consists of red blood cells, white blood cells, and platelets, all suspended within plasma, the liquid, noncellular part of the blood. In humans and other vertebrates, red blood cells contain the respiratory pigment that transports oxygen. White blood cells defend the body against disease-causing microorganisms (see Chapter 45). Platelets, small fragments broken off from large cells in the bone marrow, play a key role in blood clotting. Plasma consists of water, proteins, salts, and a variety of soluble chemical messengers such as hormones that it transports from one part of the body to another. We discuss blood in Chapter 44.

Muscle tissue is specialized to contract Most animals move by contracting the long, cylindrical or spindleshaped cells of muscle tissue. Muscle cells are called muscle fibers because of their length. Each muscle fiber contains many thin, longitudinal, parallel contractile units called myofibrils. Two proteins, myosin and actin, are the chief components of myofibrils. Myosin and actin play a key role in contraction of muscle fibers. Many invertebrates have skeletal and smooth muscle. Vertebrates have three types of muscle tissue: skeletal, cardiac, and smooth (TABLE 39-3 on page 830). Skeletal muscle makes up the large muscle masses attached to the bones of the body. Skeletal muscle fibers are very long, and each fiber has many nuclei. The nuclei of skeletal muscle fibers are also unusual in their position. They lie just under the plasma membrane, which frees the entire central part of the skeletal muscle fiber for the myofibrils. This adaptation appears to increase the efficiency of contraction. When skeletal muscles contract, they move parts of the body. Whereas skeletal muscle fibers are generally under voluntary control, you do not normally contract your cardiac and smooth muscle fibers at will. Light microscopy shows that both skeletal and cardiac fibers have alternating light and dark transverse stripes, or striations, that change their relative sizes during contraction. Striated muscle fibers contract rapidly but cannot remain contracted for a long period. They must relax and rest momentarily before contracting again. (Muscle contraction is discussed in Chapter 40.) Cardiac muscle is the main tissue of the heart. When cardiac muscle contracts, the heart pumps the blood. The fibers of cardiac muscle join end to end, and they branch and rejoin to form complex networks. One or two nuclei lie within each fiber. A characteristic feature of cardiac muscle tissue is the presence of intercalated discs, specialized junctions where the fibers join. Smooth muscle occurs in the walls of the digestive tract, uterus, blood vessels, and many other internal organs. Contraction of smooth muscle is necessary for these organs to perform certain functions. For example, smooth muscle contraction in the wall of the digestive tract moves food through the digestive tract. When smooth muscle in the walls of arterioles (small arteries) contracts,

(a) The human skeleton consists mainly of bone.

(b) A bone cut open to expose its internal structure.

Spongy bone

Cytoplasmic extensions

Lacuna

Compact bone

Haversian canal

Blood vessel

(c) Blood vessels and nerves run through the Haversian canal within each osteon of compact bone.

Osteon

Matrix Cytoplasmic extensions

Osteocyte

(d) The bone matrix is rigid and hard. Osteocytes become trapped within lacunae but communicate with one another by way of cytoplasmic extensions that extend through tiny canals.

FIGURE 39-2 Animated Bone

the blood vessels constrict, raising blood pressure. Each spindleshaped smooth muscle fiber contains a single, central nucleus.

Nervous tissue controls muscles and glands Nervous tissue consists of neurons and glial cells. Neurons are specialized for receiving and transmitting signals. Glial cells sup-

port and nourish the neurons, destroy pathogens, and modulate transmission of impulses (FIG. 39-3). A typical neuron has a cell body containing the nucleus as well as two types of cytoplasmic extensions (discussed in Chapter 41). Dendrites are cytoplasmic extensions specialized for receiving signals and transmitting them to the cell body. The single axon transmits signals, called nerve impulses, away from the cell body.

TABLE 39-3

Muscle Tissues

Location Type of control Shape of fibers Striations Number of nuclei per fiber Position of nuclei Speed of contraction Resistance to fatigue (with repetitive contraction)

Skeletal

Cardiac

Smooth

Attached to skeleton Voluntary Elongated, cylindrical, blunt ends Present Many Peripheral Most rapid Least

Walls of heart Involuntary Elongated, cylindrical, fibers that branch and fuse Present One or two Central Intermediate (varies) Intermediate

Walls of stomach, intestines, etc. Involuntary Elongated, spindle shaped, pointed ends Absent One Central Slowest Greatest

Nuclei

Striations

Nuclei

Nuclei

Intercalated discs

Skeletal muscle fibers

Cardiac muscle fibers

Smooth muscle fibers

In this chapter, we have focused on normal tissues. For a discussion of some abnormal tissues, see Inquiring About: Unwelcome Tissues: Cancers.

Neurons

Ed Reschke

Dendrite Nuclei of glial cells

Axon

100 μm

FIGURE 39-3 LM of nervous tissue Neurons transmit information in the form of electrochemical signals. Glial cells support, protect, and nourish neurons. They also communicate and help regulate neural function.

Axons range in length from 1 or 2 mm to more than a meter. Those extending from the spinal cord down the arm or leg in a human, for example, may be a meter or more in length. Certain neurons receive signals from the external or internal environment and transmit them to the spinal cord and brain. Other neurons relay, process, or store information. Still others transmit signals from the brain and spinal cord to the muscles and glands. Neurons communicate at junctions called synapses. A nerve consists of a great many neurons bound together by connective tissue.

Tissues and organs make up the organ systems of the body Tissues associate to form organs. Although an animal organ may be composed mainly of one type of tissue, other types are needed to support, protect, provide a blood supply, and transmit information. For example, the heart is mainly cardiac muscle tissue, but its chambers are lined with endothelium and its walls contain blood vessels made of endothelium, smooth muscle, and connective tissue. The heart also has nerves that transmit information and help regulate the rate and strength of its contractions. An organized group of tissues and organs that together perform a specialized set of functions make up an organ system. Working together in a very coordinated way, organ systems perform the functions required by the organism. We can identify 11 major organ systems that work together to carry out the physiological processes of a mammal: integumentary, skeletal, muscular, digestive, cardiovascular, immune (lymphatic), respiratory, urinary, nervous, endocrine, and reproductive systems. FIGURE 39-4 summarizes the principal organs and functions of each organ system.

Review ■

■

■

■

What are the main differences in structure and function between epithelial tissue and connective tissue? What type of tissue lines the air sacs of the lungs? How is its structure adapted to its function? What are some differences between the three types of muscle tissue? How is the structure of each type adapted for its function? What are the main functions of each of the following organ systems? (1) respiratory, (2) urinary, and (3) endocrine? (Consult Fig. 39-4 for help.)

In q u ir in g A b o u t U N W E LCO M E T I S S U E S : C A N C E R S through these walls and are transported to new sites. Worldwide, cancer causes more than seven million deaths each year. In the United States, cancer is the second leading cause of death. One in three people in the United States develops cancer at some time in his or her life. Currently, the key to survival is early diagnosis and treatment with some combination of surgery, hormonal treatment, radiation therapy, chemotherapy, inhibitors that suppress development of new blood vessels, immunotherapy, and targeted therapies. Many new treatments are under investigation, including agents that inhibit the development of new blood vessels. Immunotherapy for cancer will be discussed in Chapter 45. Cancer is a large family of closely related diseases (there are hundreds of distinct varieties), and treatment must be tailored to the particular type of cancer. Alleles of some genes appear to affect an individual’s level of tolerance to carcinogens. Most cancers are thought to be triggered by carcinogens, cancerproducing agents, in the environment and by diet and lifestyle. You can decrease risk of developing cancer by following these recommendations: 1. Do not smoke or use tobacco. Smoking is responsible for more than 80% of lung

Moredum Animal Health LTD /Photo Researchers, Inc.

A neoplasm (“new growth”) is an abnormal mass of cells. A tumor is a neoplasm that has formed a lump. A neoplasm may be benign or malignant (cancerous). A benign (“kind”) neoplasm tends to grow slowly, and its cells stay together. Because benign tumors form masses with distinct borders, they can usually be removed surgically. A malignant (“wicked”) neoplasm, or cancer, usually grows much more rapidly and invasively than a benign tumor. In Chapter 17 you learned that cancer results from abnormal expression of specific genes critical for cell division (see Fig. 17-20). Most cancer cells divide in an uncontrolled way. Unlike normal cells, which respect one another’s boundaries and form tissues in an orderly, organized manner, cancer cells grow helter-skelter and infiltrate normal tissues. They apparently no longer receive or respond appropriately to signals from surrounding cells; communication is lacking (see figure). When a cancer cell multiplies, all the cells derived from it are also abnormal. Unlike the cells of benign neoplasms, cancer cells do not retain normal structural features. Cancers that develop from connective tissues or muscle are referred to as sarcomas. Most human cancers originate in epithelial tissue and are called carcinomas. This group includes breast, prostate, colon, lung cancer, and most ovarian cancers. Death from cancer results from the effects of metastasis, migration of cancer cells through blood or lymph channels to other parts of the body. Once there, cancer cells multiply, forming new malignant neoplasms that interfere with the normal functions of the tissues being invaded. Cancer often spreads so rapidly and extensively that surgeons cannot locate or remove all the malignant masses. Solid tumors, which account for more than 85% of cancer deaths, require blood vessels to ensure delivery of nourishment and oxygen. Some neoplasms grow to several millimeters in diameter and then enter a dormant stage, which may last for months or even years. Eventually, cancer cells release a chemical substance that stimulates nearby blood vessels to develop new capillaries. These blood vessels grow into the abnormal mass of cells. Nourished by its new blood supply, the neoplasm begins to grow rapidly. Newly formed blood vessels have leaky walls that provide a route for metastasis. Malignant cells enter the blood

cancer cases, and it increases the risk for many other cancers. 2. Avoid prolonged exposure to the sun. When in the sun, use sunscreen or sunblock. Exposure to the sun is responsible for almost all the 400,000 cases of skin cancer reported each year in the United States alone. 3. Eat a healthy diet, including fresh, unprocessed fruits, vegetables, and grains. Limit intake of red meat. Avoid smoked, salt-cured, and nitrite-cured foods. Limit intake of alcoholic beverages. Reduce intake of foods and drinks that contribute to weight gain. Obesity increases the risk of cancer. 4. Exercise. Physical inactivity has been linked with increased risk of colon, breast, and other cancers. 5. Avoid unnecessary exposure to X-rays. 6. Self-examination/screening can lead to early diagnosis. Women should examine their breasts each month, have regular mammograms, and obtain annual Papanicolaou (Pap) and human papilloma virus (HPV) tests. Certain HPV strains can cause cervical cancer; vaccines are now available. 7. Self-examination/screening. Men should regularly examine their testes and have prostate examinations yearly after age 50. They should also have the prostatespecific antigen (PSA) blood test. These routine tests detect cancer at an early, more treatable stage. 8. Self-examination/screening. Beginning at age 50, both men and women should be screened for colorectal cancer. Detection and removal of polyps (benign growths that can become malignant) can prevent cancer.

50 μm

When cancer cells multiply, they invade normal tissues and interfere with their functions. This SEM shows a mass of malignant cells (pink) in the air sac in the center. Cancer cells that have separated from the main tumor can be seen in other air sacs. Microvilli on the surface of the cancer cells give them a fuzzy appearance.

9. Genetic testing. If you have a family history of certain cancers, you may be at higher risk genetically for cancer. Genetic tests are now available to determine if you have mutations in the BRCA1 or BRCA2 genes, which increase risk for breast, ovarian, and pancreatic cancers. Mutations in several other genes indicate increased risk for colon, uterine, stomach, and urinary tract cancer. If you are at increased risk for cancer, a genetic counselor can advise you of measures you can take to reduce that risk.

Hair

Skin

Fingernails

(1) INTEGUMENTARY SYSTEM Skin, nails, hair, sweat glands. Protects body, helps regulate body temperature, receives stimuli.

(3) MUSCULAR SYSTEM Skeletal muscles that enable us to move, cardiac muscle of the heart, and smooth muscle of internal organs.

(2) SKELETAL SYSTEM Bones, cartilage, ligaments, joints. Supports and protects body, important in movement, stores calcium.

Toenails Brain Arteries Nerves

Pineal Thyroid

Spinal cord

Hypothalamus Pituitary

Heart

Parathyroids Thymus

Veins

Adrenals Pancreas (islets) Ovaries

(4) NERVOUS SYSTEM Brain, spinal cord, sense organs, and nerves. Principal regulatory system.

Testes

(5) ENDOCRINE SYSTEM Ductless glands that release hormones. Works with nervous system in regulating metabolic activities and many other functions.

FIGURE 39-4 Animated The principal organ systems of the human body

(6) CARDIOVASCULAR SYSTEM Heart, blood vessels, blood. Transports nutrients, oxygen, hormones and other substances throughout body.

Pharynx Nasal Thymus cavity Thoracic duct Pharynx Lymph node (throat)

Oral cavity Salivary glands Esophagus

Oral cavity (mouth)

Liver

Larynx (voice box)

Spleen Lungs

Stomach Gallbladder

Bronchus

Pancreas Small intestine Large intestine Rectum

Diaphragm

Lymph vessels

Anus

(7) IMMUNE/LYMPHATIC SYSTEM Lymphatic vessels, lymph, lymph nodes, spleen, thymus, tonsils. Returns excess tissue fluid to blood, absorbs lipids, defends against disease.

(9) DIGESTIVE SYSTEM Digestive tract and glands that secrete digestive juices. Processes food, eliminates wastes.

(8) RESPIRATORY SYSTEM Lungs and air passageways. Supplies oxygen to blood and excretes carbon dioxide.

Kidney Ureter

Prostate gland

Urinary bladder

Vas deferens Penis

Urethra Oviduct

(10) URINARY SYSTEM Kidneys, urinary bladder, and associated ducts. Kidneys remove metabolic wastes and excess materials from blood; produce urine. Helps regulate blood chemistry.

Testis

Ovary Uterus Vagina

(11) MALE AND FEMALE REPRODUCTIVE SYSTEMS Both consist of gonads and associated structures. Sexual reproduction; maintains sexual characteristics.

Many animals are conformers for certain environmental conditions. Some of their internal states vary with changes in their surroundings. For example, most marine invertebrates conform to the salinity of the surrounding sea water. Mammals are superb regulators. They have complex homeostatic mechanisms that maintain relatively constant internal conditions despite changes in the outside environment. How do homeostatic mechanisms work? Many are feedback systems, sometimes called “biofeedback systems.”

39.2 REGULATING THE INTERNAL ENVIRONMENT ■ ■ LEARNING OBJECTIVE 7

Define homeostasis, and contrast negative and positive feedback systems.

To survive and function, animals must regulate the composition of the fluids that bathe their cells. They must maintain pH and internal temperature within relatively narrow limits. The body must also maintain the appropriate concentration of nutrients, oxygen, and other gases, ions, and compounds needed for metabolism at all times. Cells, tissues, organs, and organ systems work together to maintain appropriate conditions in the body. The balanced internal environment is referred to as homeostasis. Homeostasis is a basic concept in physiology. First coined by U.S. physiologist Walter Cannon, the word homeostasis is derived from the Greek homoios, meaning “same,” and stasis, “standing.” Although, the internal environment never really stays the same, it is a dynamic equilibrium in which conditions are maintained within narrow limits, which we call a steady state. The control processes that maintain these conditions are homeostatic mechanisms. Stressors, changes in the internal or external environment that affect normal conditions within the body, continuously challenge homeostasis. An internal condition that moves out of its homeostatic range (either too high or too low) causes stress. An organism functions effectively because homeostatic mechanisms continuously operate to manage stress.

KEY POINT

Negative feedback systems restore homeostasis In a negative feedback system, a change in some steady state (for example, normal body temperature) triggers a response that counteracts, or reverses, the change. A sensor detects a change, a deviation from the normal condition, or set point. The sensor signals an integrator, or control center. Based on the input of the sensor, the integrator activates homeostatic mechanisms that restore the steady state (FIG. 39-5). The response counteracts the inappropriate change, thereby restoring the steady state. Note that in a negative feedback system, the response of the integrator is opposite (negative) to the output of the sensor. When some condition varies too far from the steady state (either too high or too low), a control system using negative feedback brings the condition back to the steady state. For example, when the glucose concentration in the blood decreases below its homeostatic level, negative feedback systems increase its concentration. Most homeostatic mechanisms in the body are negative feedback systems. Let us discuss a specific example of negative feedback systems: heat regulation. The temperature-regulating system in the human body is somewhat similar to how you regulate temperature in your home. You might set a particular room temperature on your ther-

In a negative feedback system, the response of the integrator is opposite to the input of the sensor; for example, if the glucose concentration in the blood is too low, alpha cells in the pancreas secrete a hormone that increases glucose concentration.

Stressor

HOMEOSTASIS

5

Normal condition (set point) restored.

4

Integrator activates effectors (homeostatic mechanisms).

3

FIGURE 39-5 Animated Negative feedback

Sensor signals integrator (control center).

1

2

Stressor causes deviation from set point.

Sensor detects change from set point.

Change (door opens, letting cold air in).

Stressor

HOMEOSTASIS

HOMEOSTASIS 5 5

Room temperature increases to set point.

1

Body temperature increases to set point.

4 4

Furnace (effector) increases heat output. 2

3

Body temperature decreases below set point.

1

Room temperature decreases below set point. Integrator activates homeostatic mechanisms (effectors) that increase body temperature.

Thermometer (sensor) detects change.

3

Thermometer signals thermostat (integrator).

(a) Regulation of room temperature.

2

Sensors signal temperature-regulating center in hypothalamus (integrator).

Specialized nerve cells (sensors) detect change.

(b) Regulation of body temperature.

FIGURE 39-6 Negative feedback in temperature regulation Note that the diagram in (b) is highly simplified. Figure 39-9 shows greater detail.

mostat. If the temperature in the room falls, the thermometer in the thermostat acts as a sensor that detects a change, or deviation, from the set point (FIG. 39-6a). The thermometer sends a signal to the thermostat, which then acts as an integrator, or control center. The thermostat compares the sensor’s input with the set point. The thermostat then signals the furnace, which is the effector in this system. An effector is the device, organ, or process that helps restore the steady state. The furnace increases its heat output, a corrective response that brings the room temperature back to the set point. The thermometer no longer detects a change from the set point, so the thermostat and furnace are switched off. Basically, the negative feedback system that regulates body temperature is similar, as shown in FIGURE 39-6b. When body temperature decreases below normal limits, specialized nerve cells (sensors) signal the temperature-regulating center in the hypothalamus of the brain (the integrator). The integrator activates effectors that bring the temperature back to the set point. The return to normal temperature signals the sensors, and the temperatureregulating center switches off the effectors.

A few positive feedback systems operate in the body In a positive feedback system, a change in some steady state sets off a response that intensifies (rather than reverses) the changing condition. Although some positive feedback mechanisms are beneficial, they do not maintain homeostasis. For example, a positive feedback cycle operates during the birth of a baby. As the baby’s head pushes against the cervix (lower part of uterus), a reflex action causes the uterus to contract. The contraction forces the head against the cervix again, stimulating another contraction, and the positive feedback cycle repeats again and again until the baby is born. Some positive feedback sequences, such as those that deepen circulatory shock

following severe hemorrhage, can disrupt steady states and lead to death. A simplified example is shown in FIGURE 39-7. Homeostatic mechanisms maintain the internal environment within the physiological limits that support life. As you continue your study of animal processes, you will learn many ways in which organ systems interact to maintain the steady state of the organism. Although the nervous and endocrine systems play major roles, all organ systems participate in these regulatory processes. In the next section, we discuss some specific homeostatic mechanisms that help regulate body temperature.

Review ■

What is the basic difference between negative and positive feedback systems?

39.3 THERMOREGULATION ■ ■ LEARNING OBJECTIVES 8 Compare the costs and benefits of ectothermy. 9 Compare the costs and benefits of endothermy, and describe strategies animals use to adjust to challenging temperature changes.

Many animals have elaborate homeostatic mechanisms for regulating body temperature. Some are physiological, and others are structural or behavioral. Thermoregulation is the process of maintaining body temperature within certain limits despite changes in the surrounding temperature. Animals produce heat as a byproduct of metabolic activities. Body temperature is determined by the rate at which heat is produced and the rate at which heat is lost to, or gained from, the outside environment.

KEY POINT

Positive feedback intensifies the change taking place, moving conditions further away from homeostasis; in some situations, such as hemorrhage, the results can be fatal.

Stressor: hemorrhage Homeostasis

1

4

Cardiac output decreases (heart pumps less blood).

Loss of blood causes blood pressure to decrease.

2

3

Less blood circulates to heart.

Heart function declines.

FIGURE 39-7 Positive feedback In positive feedback, the changes that occur increase the deviation from the set point. Conditions move further from the normal range. Loss of blood decreases blood pressure. Less blood reaches the heart, so heart

function decreases. The resulting decrease in cardiac output further decreases blood pressure, bringing about conditions farther from homeostasis.

The strategies for maintaining body temperature that are available to an animal may restrict the type of environment it can inhabit. Each species has an optimal environmental temperature range. Some animals, such as snowshoe hares, snowy owls, and weasels, can survive in cold arctic regions. Others, such as the Cape ground squirrel, which inhabits South Africa, are adapted to hot tropical climates. Although some animals can survive at temperature extremes, most survive only within moderate temperature ranges.

mates during the winter. Another behavioral strategy for regulating temperature is hibernation. Some insects use a combination of structural, behavioral, and physiological mechanisms to regulate body temperature. The “furry” body of the moth helps conserve body heat. When a moth prepares for flight, it contracts its flight muscles with little movement of its wings. The metabolic heat generated enables the moth to sustain the intense metabolic activity needed for flight.

Ectotherms absorb heat from their surroundings

Endotherms derive heat from metabolic processes

Ectotherms are animals that depend on the environment for their body heat. Their body temperature is determined mainly by the changing temperature of their surroundings. Most of the heat for their thermoregulation comes from the sun. You may be surprised to learn that most animals are ectotherms. A benefit of ectothermy is that very little energy is used to maintain a high metabolic rate. In fact, an ectotherm’s metabolic rate tends to change with the weather. As a result, ectotherms have a much lower daily energy expenditure than do endotherms. They survive on less food and convert more of the energy in their food to growth and reproduction. Ectothermy also has costs. One disadvantage is that daily and seasonal temperature conditions may limit activity. Many ectotherms use behavioral strategies to adjust body temperature. For example, lizards may keep warm by burrowing in the soil at night. During the day, lizards take in heat by basking in the sun, orienting their bodies to expose the maximum surface area to the sun’s rays (FIG. 39-8a). Many animals migrate to warmer cli-

Birds and mammals, as well as some species of fish (e.g., tuna and some sharks) and some insects, are endotherms: they have homeostatic mechanisms that maintain body temperature despite changes in the external temperature. The most important benefit of endothermy is the high metabolic rate, which can be as much as six times higher than that of ectotherms. Their constant body temperature allows a higher rate of enzyme activity than is possible for ectotherms living in the same habitat. Endotherms respond more rapidly to internal and external stimuli and can be active even in low winter temperatures. However, these animals must pay the high energy cost of thermoregulation during times when they are inactive. You must maintain your body temperature even when you are asleep. Endotherms have structural adaptations for maintaining body temperature. For example, the insulating feathers of birds, hair of mammals, and insulating layers of fat in birds and mammals reduce heat loss from the body. Birds and mammals also have behavioral

that signals the thyroid gland. Secretion of thyroid hormones increases, which raises metabolic rate. Body tissues increase heat production. The hypothalamus also sends neural signals that cause blood vessels in the skin to constrict. As a result, less heat is brought to the body surface. In addition, nerves signal muscles to shiver or let you move muscles voluntarily to increase body temperature.

Many animals adjust to challenging temperature changes Animals adjust to seasonal changes, a process called acclimatization. A familiar example is the thickening of a dog’s coat in the winter. As water temperature decreases during fall and winter, a trout’s enzyme systems decrease their level of activity, allowing the trout to remain active at the lowest metabolic cost. When stressed by cold, many small endotherms sink into torpor, a short-term state in which metabolic rate decreases, sometimes dramatically. Torpor saves the energy that the animal would use to maintain a high body temperature. Instead, body temperature decreases below normal levels. The decrease in body temperature is brought about by a decrease in the temperature set point in the hypothalamus. Heart and respiratory rates decrease, and animals are less responsive to external stimulation. Daily torpor occurs in some small endotherms, for example, hummingbirds and shrews. We can think of torpor as an adaptive hypothermia that helps animals survive during times of cold temperatures.

(a) Ectotherm. The rainbow agama (Agama agama) increases its body temperature by sunning itself. This insect-eating lizard was photographed in Serengeti National Park, Tanzania.

FIGURE 39-8 Behavioral adaptations for thermoregulation

Frans Lanting/Minden Pictures

Richard J. Green /Photo Researchers, Inc.

adaptations for maintaining body temperature. The Cape ground squirrel positions its tail to shade its body from the direct rays of the sun. Elephants spray themselves with cool water. Endotherms have a variety of physiological mechanisms for maintaining temperature homeostasis. They regulate heat production and regulate heat exchange with the environment. Most of their body heat comes from their own metabolic processes (see Inquiring About: Electron Transport and Heat, in Chapter 8). In mammals, receptors located in the hypothalamus of the brain and in the spinal cord regulate temperature. Heat from metabolic activities can be increased directly or can be increased indirectly by the action of hormones (such as thyroid hormones) that increase metabolic rate. Heat production is increased by contracting muscles, and in cold weather many animals shiver. When body temperature rises, birds and many mammals pant and some mammals sweat (FIG. 39-8b). These processes provide fluid for evaporation, the conversion of a liquid, such as sweat, to water vapor. (Recall from Chapter 2 that when molecules enter the vapor phase, they take their heat energy with them.) Heat transfers from the body to the surroundings, resulting in evaporative cooling. In the human body, about 2.5 million sweat glands secrete sweat. As sweat evaporates from the skin surface, body temperature decreases. Constriction and dilation of capillaries in the skin are also homeostatic mechanisms for regulating body temperature. In humans, when body temperature increases above normal, specialized nerve cells signal the temperature-regulating center in the hypothalamus (FIG. 39-9). This center sends messages, by way of nerves to the sweat glands, which increase sweat secretion. At the same time, the hypothalamus sends messages to smooth muscle in the walls of blood vessels in the skin that cause them to dilate. More blood circulates through the skin, bringing heat to the body surface. The skin acts as a heat radiator that allows heat to radiate from the body surface into the environment. These homeostatic mechanisms help return body temperature to normal. When body temperature decreases below normal, the hypothalamus signals the anterior pituitary gland to release a hormone

(b) Endotherm. The body temperature of this baby sooty tern (Sterna fuscata ) is reduced as heat leaves the body through its open mouth. Photographed in Hawaii.

KEY POINT

The hypothalamus maintains homeostasis of body temperature by activating mechanisms that cool (upper arrows) or warm (lower arrows) the body.

Decreased muscle activity Nerves

Increased sweating/ panting

Evaporation

Smooth muscle in skin blood vessels relaxes.

Blood vessels dilate

Sensors signal temperature-regulating center in hypothalamus (integrator).

Specialized nerve cells (sensors) detect change from set point.

Body temperature increases

Stressors

Body temperature decreases (normal condition restored). Stressors

HOMEOSTASIS

Body temperature increases (normal condition restored).

Body temperature decreases

Specialized nerve cells (sensors) detect change from set point. Sensors signal temperature-regulating center in hypothalamus (integrator). Blood vessels constrict.

Increase in voluntary movement; shivering Nerves

Increase in metabolic rate

Smooth muscle in skin blood vessels contracts. Anterior pituitary gland

Thyroid gland

FIGURE 39-9 Animated Regulation of temperature in the human body

Hibernation is long-term torpor in response to winter cold and scarcity of food. Animals that hibernate store unsaturated fats as energy sources. Estivation is a state of torpor caused by lack of food or water during periods of high temperature. During estivation some mammals retreat to their burrows. The cactus mouse enters a state of hibernation during the winter in response to cold and scarcity of food. In summer, it estivates in response to lack of either food or water.

Review ■

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How do ectotherms adjust body temperature? What are some costs and benefits of ectothermy? What are some costs and benefits of endothermy? How is body temperature regulated in humans? Draw a diagram to illustrate your answer. How do hibernation and estivation differ?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

39.1 (page 822) 1 Compare the structure and function of the four main kinds of animal tissues: epithelial, connective, muscle, and nervous tissues. A tissue consists of a group of similarly specialized cells that associ■ ate to perform specific functions. Epithelial tissue (epithelium) forms a continuous layer, or sheet, of cells covering a body surface or lining a body cavity. Epithelial tissue functions in protection, absorption, secretion, and sensation. Connective tissue consists of relatively few cells separated by an ■ intercellular substance, composed of fibers scattered throughout a matrix. The intercellular substance contains three types of fibers. Connective tissue contains specialized cells such as fibroblasts and ■ macrophages. Connective tissue joins other tissues of the body, supports the body and its organs, and protects underlying organs. Muscle tissue consists of cells specialized to contract. Each cell is ■ an elongated muscle fiber containing many contractile units called myofibrils. Nervous tissue consists of elongated cells called neurons, specialized for transmitting impulses, and glial cells, which support and nourish the neurons. 2 Compare the main types of epithelial tissue and describe their functions. Epithelial cells may be squamous, cuboidal, or columnar in shape. ■ Epithelial tissue may be simple, stratified, or pseudostratified (summarized in Table 39-1). Simple squamous epithelium lines blood vessels and the air sacs ■ in the lungs; it permits exchange of materials by diffusion. Simple cuboidal and columnar epithelia line passageways and are specialized for secretion and absorption. Stratified squamous epithelium forms the outer layer of the skin and lines passageways into the body; it provides protection. Pseudostratified epithelium also lines passageways and protects underlying tissues. Some epithelial tissue is specialized to form glands. Goblet cells ■ are unicellular glands that secrete mucus. Goblet cells are exocrine glands that secrete their product through a duct onto an exposed epithelial surface. In contrast, endocrine glands release hormones into the interstitial fluid or blood. An epithelial membrane consists of a sheet of epithelial tissue and ■ a layer of underlying connective tissue. A mucous membrane lines a cavity that opens to the outside of the body. A serous membrane lines a body cavity that does not open to the outside.

3 Compare the main types of connective tissue and describe their functions. ■

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The cells of connective tissue are embedded in an intercellular substance that consists of microscopic collagen fibers, elastic fibers, and reticular fibers (thin branched fibers) scattered through a matrix, a thin gel of polysaccharides. Loose connective tissue consists of fibers running in various directions through a semifluid matrix; it forms a covering for nerves, blood vessels, and muscles. Dense connective tissue is strong but less flexible than loose connective tissue. It forms tendons, cords that connect muscles to bones, and ligaments, cables that connect bones to one another. Elastic connective tissue consists of bundles of parallel elastic fibers; it is found in lung tissue and in walls of large arteries. Reticular connective tissue, which consists of interlacing reticular fibers, forms a supporting framework for many organs. Adipose tissue is made up of fat cells; it is found along with loose connective tissue in subcutaneous tissue. Cartilage consists of chondrocytes that lie in lacunae, small cavities in a hard matrix. Osteocytes secrete and maintain the matrix of bone. Unlike cartilage, bone is quite vascular. Cartilage and bone form the skeleton of vertebrates. Blood and lymph are circulating tissues that help various parts of an animal communicate with one another. The intercellular substances of blood and lymph are more fluid than those of other tissues.

4 Contrast the three types of muscle tissue and describe their functions. Skeletal muscle is striated and under voluntary control. Each elongated, cylindrical muscle fiber has several nuclei. When skeletal muscles contract, they move parts of the body. Cardiac muscle is also striated, but its contractions are involuntary. ■ Its elongated, cylindrical fibers branch and fuse; each fiber has one or two central nuclei. When this muscle contracts, the heart pumps the blood. Smooth muscle contracts involuntarily; its elongated, spindle■ shaped fibers lack striations. Each fiber has a single central nucleus. Smooth muscle is responsible for movement of body organs; for example, it pushes food through the digestive tract. 5 Relate the structure of the neuron to its function. The elongated neuron is adapted for receiving and transmitting in■ formation. Dendrites receive signals and transmit them to the cell body. The axon transmits signals away from the cell body to other neurons or to a muscle or gland. A synapse is a junction between neurons. 6 Briefly describe the organ systems of a mammal and summarize the functions of each organ system. Tissues and organs work together, forming organ systems. In ■ mammals, 11 organ systems work together to carry out the functions required by the organism. These include the integumentary, skeletal, muscular, nervous, endocrine, cardiovascular, immune (lymphatic), respiratory, digestive, urinary, and reproductive systems (summarized in Figure 39-4). Each organ system functions to maintain homeostasis. ■

Watch body systems work together by clicking on the figure in CengageNOW.

39.2 (page 834) 7 Define homeostasis, and contrast negative and positive feedback systems. Homeostasis is the balanced internal environment, or steady state. ■ The control processes that maintain these conditions are homeostatic mechanisms, mainly negative feedback systems. When a stressor causes a change in some steady state, a response is triggered that counteracts the change. A sensor detects a change, a deviation from the desired condition, or set point. The sensor signals an integrator, or control center. Based on the output of the sensor, the integrator activates one or more effectors, organs or processes that restore the steady state.

Stressor

HOMEOSTASIS

5

Body temperature increases to set point.

4

Integrator activates homeostatic mechanisms (effectors) that increase body temperature. 3

Body temperature decreases below set point.

1

2

Sensors signal temperature-regulating center in hypothalamus (integrator).

Specialized nerve cells (sensors) detect change.

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In a positive feedback system, a deviation from the steady state sets off a series of changes that intensify (rather than reverse) the changes.

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39.3 (page 835) 8 Compare the costs and benefits of ectothermy. Animals have structural, behavioral, and physiological strategies for thermoregulation, the process of maintaining body temperature within certain limits despite changes in the surrounding temperature. ■ In ectotherms, body temperature depends to a large extent on the temperature of the environment. Many ectotherms use behavioral strategies to adjust body temperatures. A benefit of ectothermy is that very little energy is used to maintain the metabolic rate. A disadvantage is that activity may be limited by daily and seasonal temperature conditions. 9 Compare the costs and benefits of endothermy, and describe strategies animals use to adjust to challenging temperature changes. ■

■ ■

Endotherms have homeostatic mechanisms for regulating body temperature within a narrow range. The most important benefit of endothermy is a high metabolic rate. Another advantage is that constant body temperature allows a higher rate of enzyme activity. Many endotherms are active even in low winter temperatures. A disadvantage of endothermy is its high energy cost. Acclimatization is the process of adjustment to seasonal changes. When challenged by a drop in surrounding temperature, many small endotherms enter a state of torpor. Torpor is an adaptive hypothermia; body temperature is maintained below normal levels. Hibernation is long-term torpor in response to winter cold. Estivation is torpor caused by lack of food or water during summer heat. Explore negative feedback and temperature regulation in humans and other animals by clicking on the figures in CengageNOW.

T E S T YO U R U N D E R S TA N D I N G 1. A group of closely associated cells that carry out specific functions is (a) a colony (b) a tissue (c) an organ system (d) an effector (e) a homeostatic mechanism 2. Epithelial cells that produce and secrete a product into a duct form (a) an endocrine gland (b) an exocrine gland (c) a pseudostratified membrane (d) an endocrine or exocrine gland (e) a serous membrane 3. A serous membrane (a) lines a body cavity that does not open to the outside of the body (b) consists of loose connective tissue (c) has a framework of reticular connective tissue (d) covers the body (e) lines the digestive and respiratory systems 4. Tissue that contains fibroblasts and a great deal of intercellular substance is (a) connective tissue (b) muscle tissue (c) nervous tissue (d) pseudostratified epithelium (e) simple squamous epithelium

11. Which of the following is not true of torpor? (a) it is an adaptive state of low body temperature (b) the temperature set point in the hypothalamus decreases (c) metabolic rate increases (d) it is a strategy used by some endotherms (e) respiratory rate decreases 12. An ectotherm (a) has a higher rate of enzyme activity than a typical endotherm (b) has a variety of homeostatic mechanisms for regulating body temperature (c) depends on sensors in the hypothalamus to regulate temperature (d) may use behavioral strategies to help adjust body temperature (e) must expend more energy on thermoregulation than an endotherm 13. Identify and label the tissues. See Tables 39-1 and 39-2 to check your answers.

5. Dense connective tissue is likely to be found in (a) the outer layer of skin (b) the framework of the lymph nodes (c) bone (d) the air sacs of lungs (e) tendons 6. Tissue that contracts and is striated and involuntary is (a) connective tissue (b) smooth muscle (c) skeletal muscle (d) pseudostratified (e) cardiac muscle

8. The contractile elements in muscle tissue are (a) myofibrils (b) elastic fibers (c) collagen fibers (d) lacunae (e) dendrites 9. Which system has the homeostatic function of helping regulate volume and composition of blood and body fluids? (a) integumentary (b) muscular (c) reproductive (d) urinary (e) exocrine 10. Many homeostatic functions are maintained by (a) negative feedback systems (b) positive feedback systems (c) set points (d) stressors (e) exocrine glands

Ed Reschke/Peter Arnold, Inc.

7. Glial cells are characteristic of (a) muscle that is striated and voluntary (b) cardiac muscle tissue (c) nervous tissue (d) stratified epithelium (e) loose connective tissue

Ed Reschke/Peter Arnold, Inc.

Ed Reschke/Peter Arnold, Inc.

CRITICAL THINKING 1. Imagine that all the epithelium in an animal, such as a human, suddenly disappeared. What effects might this have on the body and its ability to function? 2. What would connective tissue be like if it had no intercellular substance? What effect would the absence of intercellular substance have on the body? 3. A high concentration of carbon dioxide in the blood and interstitial fluid results in more rapid breathing. Explain this observation in terms of homeostasis. 4. EVOLUTION LINK. From an evolutionary perspective, why do you think most animals are ectotherms? (Hint: What are some benefits of ectothermy?)

5. SCIENCE, TECHNOLOGY, AND SOCIETY. Imagine that you are a healthcare professional. If a patient told you she had a family history of breast cancer, would you advise her to be genetically tested? Why or why not? What if you had a family history of cancer. Would you want to be tested? Additional questions are available in CengageNOW at www.cengage.com/ login.

40

Protection, Support, and Movement

Stephen Dalton/NHPA

Grasshopper in midjump. The grasshopper (Tropidacris dux) uses powerful muscles in its hind legs to catapult itself into the air. Insect muscles attach to the exoskeleton.

KEY CONCEPTS

I

n Chapter 39 we described animal tissues, organs, and organ systems, laying the foundation for the more detailed discussions in this and later

40.1 Many structures and processes have evolved in

chapters. Here we focus on epithelial coverings, skeletons, and muscles—

animals for protection, support, and movement. Epithelial coverings are adapted for protecting underlying tissues and may be specialized for gas exchange, sensory reception, secretion, or other functions.

structures that are closely interrelated in the functions of protection,

40.2 Skeletal systems, whether they are hydrostatic

support, and movement. In our survey of the animal kingdom (Chapters 30 to 32), we described many adaptations involving these structures. For example, we discussed how the arthropod exoskeleton contributes to

skeletons, exoskeletons, or endoskeletons, are adapted for supporting and protecting the body and for transmitting mechanical forces important in movement.

the great biological success of insects. In this chapter we compare these

40.3 Muscles are adapted for contracting; during contrac-

The epithelial coverings of animals protect underlying tissues and

tion, energy from ATP is used to slide muscle filaments so that the muscle shortens. Body parts move when muscles pull on them.

systems in several animal groups and then focus on their structure and function in mammals. have several other functions as well. One of the most interesting is secretion of peptides and other substances that kill harmful bacteria and certain other pathogens. Frogs and toads, in particular, secrete a variety of potent peptides that protect them against harmful bacteria, fungi, and protozoa. Secretions of the frog Rana ridibanda have been shown to kill “superbugs,” such as MRSA (methicillin-resistant Staphylococcus aureus; see Chapter 25). The skin of amphibians is a promising potential source of new antibiotics and other drugs. Most animals are also protected by a skeletal system. Whether it is a fluid-filled compartment, a shell, or a system of bones, the skeletal sys-

tem protects internal organs and provides support for the body. The skeletal system and muscular system work together to produce movement. Muscle tissue is specialized to contract. Although a few animals remain rooted to one spot, sweeping their surroundings with tentacles, locomotion—movement from place to place in the environment—is characteristic of most animals. Muscles responsible for locomotion anchor to the skeleton, which gives them something firm on which to act. In most vertebrates, bones serve as levers that transmit the force necessary to move various parts of the body. During the course of evolution, most animals have developed adaptations for dependable, rapid, and responsive movement. Animals creep, walk, run, jump, swim, or fly (see photograph). These adaptations enable them to find food and escape from predators. In complex animals, effective movement depends on the cooperation and interactions of several body systems, including the muscular, skeletal, nervous, circulatory, respiratory, and endocrine systems. The precisely regulated interactions among tissues, organs, and organ systems that make complex function and maintenance of homeostasis possible are a dynamic example of systems biology in action.

40.1 EPITHELIAL COVERINGS ■ ■ LEARNING OBJECTIVES 1 2

Compare the functions of the epithelial coverings of invertebrates and vertebrates. Relate the structure of vertebrate skin to its functions.

Epithelial tissue covers all external and internal surfaces of the animal body. The structure and functions of the external epithelial covering are adapted to the animal’s environment and lifestyle. In both invertebrates and vertebrates, epithelial coverings protect the body. Epithelial coverings may also be specialized for secretion, gas exchange, or temperature regulation, and typically contain receptors that receive sensory signals from the environment.

Invertebrate epithelium may secrete a cuticle In many invertebrates, the outer epithelium is specialized to secrete protective or supportive layers of nonliving material. In insects and many other animals, the secreted material forms an outer covering called a cuticle. Some animals, including corals and mollusks, secrete a shell made mainly of calcium carbonate. In many species, the epithelium contains cells that secrete lubricants or adhesives. Such cells may release odorous secretions used for communication or for marking trails. In other species, secretory cells produce toxins used for offense or defense. The earthworm epithelium secretes a lubricating mucus that promotes efficient diffusion of gases across the body wall and reduces friction during movement through the soil. Epithelial cells may be modified as sensory cells that are selectively sensitive to light; chemical stimuli; or mechanical stimuli, such as contact with an object, or pressure.

Vertebrate skin functions in protection and temperature regulation The integumentary system of vertebrates includes the skin and structures that develop from it. In many fishes, in some reptiles, and in the anteating pangolin (a mammal; see Fig. 18-15c), skin has developed into a set of scales formidable enough to be considered armor. Even human skin has considerable strength. The thin, moist skin of many amphibians is adapted for gas exchange, as well as for defense. Derivatives of skin differ considerably among vertebrates. Fishes have bony or toothlike scales. Amphibians have naked skin covered with mucus, and some species are equipped with poison glands. Reptiles have epidermal scales, mammals have hair, and birds have feathers that provide even more effective insulation than fur. The feathers of birds and the fur of mammals are insulating structures that help maintain body temperature by trapping heat in the body. Skin and its derivatives are often brilliantly colored in connection with courtship rituals, territorial displays, and other kinds of communication. In mammals, structures derived from skin include claws (modified as fingernails and toenails in primates), hair, sweat glands, oil (sebaceous) glands, and several types of sensory receptors that give mammals the ability to feel pressure, temperature, and pain. The skin of mammals also contains mammary glands, specialized in females for secretion of milk. In humans and other mammals, oil glands empty via short ducts into hair follicles. A hair follicle is the part of a hair below the skin surface, together with the surrounding epithelial tissue. Oil glands secrete sebum, a mixture of fats and waxes that inhibits the growth of harmful bacteria. Although the skin of fishes, frogs, snakes, pelicans, and elephants appears very different, the basic structure of skin is the same in all vertebrates. The outer layer of skin, the epidermis, is a waterproof protective barrier. The epidermis consists of several strata, or sublayers. The deepest is stratum basale, and the most superficial is stratum corneum (FIG. 40-1). Pigment cells in stratum basale and in the dermis below produce melanin, a pigment that contributes to the color of the skin. In stratum basale, cells divide and are pushed outward as other cells are produced below them. Epidermal cells mature as they move toward the skin surface. As they move toward the body surface, epidermal cells manufacture keratin, an elaborately coiled protein that gives the skin considerable mechanical strength and flexibility. Keratin is insoluble and serves as a diffusion barrier for the body surface. As epidermal cells move through stratum corneum, they die. When they reach the outer surface of the skin, they wear off and must be continuously replaced. Beneath the epidermis lies the dermis, a dense, fibrous connective tissue made up mainly of collagen fibers (see Fig. 40-1). Collagen imparts strength and flexibility to the skin. The dermis also contains blood vessels that nourish the skin and sensory receptors for touch, pain, and temperature. Hair follicles and, in most mammals, sweat glands are embedded in the dermis. In birds and mammals, the dermis rests on a layer of subcutaneous tissue composed mainly of adipose tissue that insulates the body from outside temperature extremes.

Openings of sweat glands Capillary

Epidermis

Stratum corneum

Nerve endings

Stratum basale

Pigment cell (melanocyte) Hair erector muscle

Dermis

Hair shaft Sensory receptor

Subcutaneous tissue Artery Vein

Hair follicle Adipose tissue Sebaceous gland

Sweat gland

FIGURE 40-1 Animated Human integumentary system The integumentary system of humans and other mammals is a complex organ system consisting of skin and structures such as hair and nails that are derived from the skin. Stratum basale consists of a single layer of dividing epithelial cells.

In humans, exposure to ultraviolet (UV) radiation, the short, invisible rays from the sun, causes the epidermis to thicken and stimulates pigment cells in the skin to produce melanin at an increased rate. Melanin is an important protective screen against the sun because it absorbs some of the harmful UV rays. An increase in melanin causes the skin to darken. The suntan so prized by sun worshippers is actually a sign that the skin has been exposed to too much UV radiation. When the melanin cannot absorb all the UV rays, the skin becomes inflamed, or sunburned. Dark-skinned people have more melanin and so suffer less sunburn, wrinkling, and skin cancer but still require protection from UV rays. UV radiation damages DNA, causing mutations that lead to malignant transformation of cells (see Chapter 17). Skin cancer is the most common type of cancer in humans, and most cases of skin cancer are caused by excessive, chronic exposure to UV radiation. The incidence of malignant melanoma, the most lethal type of skin cancer, is increasing faster than any other type of cancer. Worldwide the number of cases has doubled during the past 20 years. Greater understanding of the Ras/ERK cell signaling pathway has led to research aimed at developing targeted drugs for treating melanoma. (Recall from Chapter 6 that this pathway is the main signaling cascade for cell division and differentiation.)

Review ■ ■ ■ ■

How are the body coverings of invertebrates and vertebrates alike? How is the structure of vertebrate skin related to its specific functions? What properties does keratin give human skin? What is the function of melanin?

40.2 SKELETAL SYSTEMS ■ ■ LEARNING OBJECTIVES 3 4 5

Compare the structure and functions of different types of skeletal systems, including the hydrostatic skeleton, exoskeleton, and endoskeleton. Describe the main divisions of the vertebrate skeleton and the bones that make up each division. Describe the structure of a typical long bone and differentiate between endochondral and intramembranous bone development.

6 Compare the main types of vertebrate joints.

In addition to having an epithelial covering, many animals are protected by a skeletal system that forms the framework of the body. The skeleton also functions in locomotion. Muscles typically act on hard structures such as chitin or bone. These skeletal structures transmit and transform mechanical forces generated by muscle contraction into the variety of motions that animals make. Most invertebrates have hydrostatic skeletons or exoskeletons. (You may recall from Chapter 32 that echinoderms are an exception.) Vertebrates have endoskeletons of cartilage or bone.

In hydrostatic skeletons, body fluids transmit force Many soft-bodied invertebrates have hydrostatic skeletons made of fluid-filled body compartments. Imagine an elongated balloon full of water. If you pull on the balloon, it lengthens and becomes thinner. It does the same if you squeeze it. Conversely, if you push

the ends toward the center, the balloon shortens and thickens. Many invertebrates, including most cnidarians, flatworms, annelids, and roundworms, have hydrostatic skeletons that work something like a balloon filled with water. Fluid in a closed compartment of the body is held under pressure. When muscles in the compartment wall contract, they push against the tube of fluid. Because fluids cannot be compressed, the force is transmitted through the fluid, changing the shape and movement of the body. In Hydra and other cnidarians, cells of the two body layers can contract. The contractile cells in the outer epidermal layer lie longitudinally, whereas the contractile cells of the inner layer (the gastrodermis) are arranged circularly around the central body axis (FIG. 40-2). The two groups of cells work in antagonistic fashion: what one can do, the other can undo. When the epidermal (longitudinal) layer contracts, the hydra shortens. Due to the fluid in the gastrovascular cavity, force is transmitted so that the hydra thickens as well. In contrast, when the inner (circular) layer contracts, the hydra thins and its fluid contents force it to lengthen. Mechanically, the hydra is a bag of fluid. The fluid acts as a hydrostatic skeleton because it transmits force when the contractile cells contract against it. (Although technically not a closed compartment, the gastrovascular cavity functions as a hydrostatic skeleton because its opening is small.) Hydrostatic skeletons permit only crude mass movements of the body or its appendages. Delicate movements are difficult because force tends to be transmitted equally in all directions throughout the entire fluid-filled body of the animal. For example, it is not easy for the hydra to thicken one part of its body while thinning another.

Longitudinal contractile fibers of epidermal layer

Circular contractile fibers of gastrodermis

The more sophisticated hydrostatic skeleton of the annelid worm lets it move with more flexibility. An earthworm’s body consists of a series of segments divided by transverse partitions, or septa (see Fig. 31-19). The septa isolate portions of the body cavity and the coelomic fluid they contain. This arrangement allows the hydrostatic skeletons of each segment to be largely independent of one another. Thus, contraction of the circular muscle in the elongating anterior end does not interfere with the action of the longitudinal muscle in the segments at the posterior end. You can find some examples of hydrostatic skeletons in invertebrates equipped with shells or endoskeletons, and even in vertebrates with endoskeletons of cartilage or bone. For example, sea stars and sea urchins have an endoskeleton, but they move their tube feet by an ingenious adaptation of a hydrostatic skeleton (see Chapter 32). And the human penis becomes erect and stiff because of the turgidity of pressurized blood in its internal spaces.

Mollusks and arthropods have nonliving exoskeletons In most animals, the skeleton is a lifeless shell, or exoskeleton, deposited atop the outer epithelial covering. In mollusks, the exoskeleton is a calcium carbonate shell secreted by the mantle, a thin sheet of epithelial tissue that extends from the body wall. The exoskeleton provides protection, a retreat used in emergencies, with the bulk of the naked, tasty body exposed at other times. Exoskeletons of arthropods serve not only to protect but also to transmit forces. In this respect they are comparable to the skeletons of vertebrates. The arthropod exoskeleton is a nonliving cuticle that contains the polysaccharide chitin. Although it is a continuous, one-piece sheath covering the entire body, the arthropod exoskeleton varies greatly in thickness and flexibility. Large, thick, inflexible plates are separated from one another by thin, flexible joints arranged segmentally. Enough joints are provided to make the arthropod’s body as flexible as those of many vertebrates. The arthropod exoskeleton is adapted to a vast variety of lifestyles, and parts of it are modified to function as specialized tools or weapons. A disadvantage of the rigid arthropod exoskeleton is that to accommodate growth, an arthropod must molt, that is, shed its exoskeleton and replace it with a new, larger one (FIG. 40-3). Recall from Chapter 31 that this process, also called ecdysis, is characteristic of the Ecdysozoa, a major branch of invertebrates. During ecdysis the animal is weak and vulnerable to predators.

Internal skeletons are capable of growth (a) Contraction of circular contractile fibers elongates the body.

(b) Contraction of longitudinal fibers shortens the body.

FIGURE 40-2 Hydrostatic skeleton In Hydra, fluid in the gastrovascular cavity transmits force when contractile cells in the body wall contract against it. The longitudinally arranged contractile cells of Hydra are antagonistic to the cells arranged circularly around the body axis.

Echinoderms and chordates have endoskeletons. This internal skeleton consists of plates or shafts of calcium-impregnated tissue (such as cartilage or bone). Composed of living tissue, the endoskeleton grows along with the animal as a whole. The echinoderm endoskeleton consists of spines and plates of calcium salts embedded in the body wall, beneath an epidermis that covers the body. This endoskeleton forms what amounts to an internal shell that provides support and protection (FIG. 40-4). Many echinoderm endoskeletons bear spines that project to the outer surface.

Patricia Jordan/Peter Arnold, Inc.

Judy Davidson/Science Photo Library/Photo Researchers, Inc.

FIGURE 40-4 The echinoderm endoskeleton As in other echinoderms, the endoskeleton of the red sea star (Formia milleporella) is composed of spines and plates of nonliving calcium salts embedded in the body wall. The endoskeleton provides support and protection.

FIGURE 40-3 Ecdysis A greengrocer cicada (Magicicada) requires 13 years to mature. It then emerges from the soil, climbs a tree, and sheds its exoskeleton, prior to reproducing.

You are probably most familiar with the endoskeleton of vertebrates. The internal vertebrate skeleton provides support and protection and transmits muscle forces. Members of class Chondrichthyes (sharks and rays) have skeletons of cartilage, but in most vertebrates the skeleton consists mainly of bone. Many bones form systems of levers that transmit muscle forces. Bones store calcium and are important in maintaining homeostatic levels of calcium in the blood.

The vertebrate skeleton has two main divisions The two main divisions of the vertebrate skeleton are the axial and appendicular skeletons (FIG. 40-5). The axial skeleton, located along the central axis of the body, consists of the skull, vertebral column, ribs, and sternum (breastbone). The appendicular skeleton consists of the bones of the limbs (arms and legs) plus the bones making up the pectoral (shoulder) girdle and most of the pelvic (hip) girdle; these girdles connect the limbs to the axial skeleton. The skull, the bony framework of the head, consists of the cranial and facial bones. In the human, 8 cranial bones enclose the brain, and 14 bones make up the facial portion of the skull. Several cranial bones that are single in the adult human result from the fusion of 2 or more bones that are separate in the fetus or newborn. The vertebrate spine, or vertebral column, supports the body and bears its weight. In humans, it consists of 24 vertebrae and 2 bones composed of fused vertebrae, the sacrum and coccyx. The

vertebral column consists of the cervical (neck) region, with 7 vertebrae; the thoracic (chest) region, with 12 vertebrae; the lumbar (back) region, with 5 vertebrae; the sacral (pelvic) region, with 5 fused vertebrae; and the coccygeal region, also composed of fused vertebrae. The rib cage is a bony “basket” formed by the sternum (breastbone), thoracic vertebrae, and, in mammals, 12 pairs of ribs. The rib cage protects the internal organs of the chest, including the heart and lungs. It also supports the chest wall, preventing it from collapsing as the diaphragm contracts with each breath. Each pair of ribs is attached dorsally to a separate vertebra. Of your 12 pairs of ribs, the first 7 are attached ventrally to the sternum; the next 3 are attached indirectly by cartilage; and the last 2, the “floating ribs,” have no attachments to the sternum. The pectoral girdle consists of two collarbones, or clavicles, and two shoulder blades, or scapulas. The pelvic girdle consists of a pair of large bones, each composed of 3 fused hipbones. Whereas the pelvic girdle is securely fused to the vertebral column, the pectoral girdle is loosely and flexibly attached to it by muscles. Each human limb consists of 30 bones and terminates in five digits, the fingers and toes. The more specialized appendages of other tetrapods may be characterized by four digits (as in the pig), three (the rhinoceros), two (the camel), or one (the horse).

A typical long bone amplifies the motion generated by muscles The radius, one of the two bones of the forearm, is a typical long bone (FIG. 40-6). Its numerous muscle attachments are arranged in such a way that the bone rotates about its long axis and operates as a lever, amplifying the motion generated by the muscles. By themselves, muscles cannot shorten enough to produce large movements of the body parts to which they are attached. Like other bones, the radius is covered by a connective tissue membrane, the periosteum, to which muscle tendons and ligaments attach. The periosteum can produce new layers of bone, thus

Bones are remodeled throughout life

Skull Clavicle

During fetal development, bones form in two ways. Long bones, such as the radius, develop from Scapula cartilage templates in a process Sternum called endochondral bone develHumerus Rib cage opment. A bone begins to ossify (become bony) in its diaphysis, Vertebrae and secondary sites of bone proPelvic Radius duction develop in the epiphyses. girdle Ulna The part of the bone between the Carpals ossified regions can grow. EventuMetacarpals ally, the ossified regions fuse. In Phalanges contrast, other bones, including the flat outer bones of the skull, Femur develop from a noncartilage, connective tissue scaffold. This process Patella is known as intramembranous bone development. Fibula Osteoblasts are bone-building Tibia cells. They secrete the protein colTarsals lagen, which forms the strong fiMetatarsals bers of bone. The compound hydroxyapatite, composed mainly of Phalanges calcium phosphate, is present in the interstitial fluid (tissue fluid). (a) Axial skeleton (brown) (b) Appendicular skeleton (brown) It automatically crystallizes around the collagen fibers to form the hard FIGURE 40-5 Animated The human skeletal system matrix of bone. As the matrix forms around the osteoblasts, they Only a few of the 206 named bones are labeled here. become isolated within the lacunae. The trapped osteoblasts are then called osteocytes. Bones are modeled during growth and remodeled continuously throughout life in response to physical stresses and other increasing the bone’s diameter. The main shaft of a long bone is its changing demands. As muscles develop in response to physical diaphysis; each expanded end is an epiphysis. In children, a disc of activity, the bones to which they are attached thicken and become cartilage, the metaphysis, lies between the epiphyses and the diastronger. As a bone grows, tissue is removed from its interior, espephysis. Metaphyses are growth centers that disappear at maturity, cially from the walls of the marrow cavity. becoming vague epiphyseal lines. Long bones have a central cavOsteoclasts are large, multinucleated cells that break down ity that contains bone marrow. Yellow marrow consists mainly of (resorb) bone. Th e osteoclasts move about, secreting hydrogen a fatty connective tissue; the red marrow in certain bones produces ions that dissolve the crystals, and enzymes that digest the colblood cells. lagen. Osteoclasts and osteoblasts are synergistic; together they The radius has a thin outer shell of compact bone, which is shape bones. Th e remodeling process is extensive—the adult huvery dense and hard. Compact bone lies primarily near the surman skeleton is completely replaced every 10 years! faces of a bone, where it provides great strength. Recall from In osteoporosis, the most common progressive, degenerative Chapter 39 that compact bone consists of interlocking spindlebone disease, bone resorption takes place more rapidly than bone shaped units called osteons (see Fig. 39-2). Within an osteon, formation. Patients lose so much bone mass that their bones beosteocytes (bone cells) lie in small cavities called lacunae (sing., come fragile, which greatly increases their risk for fracture. Most lacuna). The lacunae are arranged in concentric circles around cendrugs used to treat osteoporosis inhibit osteoclast-mediated bone tral Haversian canals. Blood vessels that nourish the bone tissue resorption. A recent addition to the treatment arsenal stimulates pass through the Haversian canals. Osteocytes are connected by osteoblasts to promote bone formation. threadlike extensions of their cytoplasm that extend through narrow channels called canaliculi. Interior to the thin shell of compact bone is a filling of spongy Joints are junctions between bones bone, which provides mechanical strength. Spongy bone consists Joints, or articulations, are junctions between two or more bones. of a network of thin strands of bone. Its spaces are filled with bone Joints facilitate flexibility and movement. At the joint, the outer marrow.

Review

Articular surface covered with cartilage Red marrow in spongy bone

■

What functions does a hydrostatic skeleton perform? How do the septa in the annelid worm contribute to the effectiveness of its hydrostatic skeleton? What are some disadvantages of an exoskeleton? some advantages?

■

What are the main bones in each division of the human skeleton?

■

How do osteoblasts and osteoclasts remodel bone?

■

Epiphysis Metaphysis

Periosteum

■

40.3 MUSCLE CONTRACTION

Yellow marrow

■ ■ LEARNING OBJECTIVES

Blood supply

7 8 9 10

Diaphysis

Compact bone

Articular cartilage

Epiphysis

FIGURE 40-6 Animated A typical long bone A long bone has a thin shell of compact bone and a filling of spongy bone that contains marrow.

surface of each bone consists of articular cartilage. One way to classify joints is according to the degree of movement they allow. The sutures between bones of the human skull are immovable joints. In a suture, bones are held together by a thin layer of dense fibrous connective tissue, which may be replaced by bone in the adult. Slightly movable joints, found between vertebrae, are made of cartilage and help absorb shock. Most joints are freely movable joints. Each is enclosed by a joint capsule of connective tissue and lined with a membrane that secretes a lubricant called synovial fluid. This viscous fluid reduces friction during movement and absorbs shock. The joint capsule is typically reinforced by ligaments, bands of fibrous connective tissue that connect bones and limit movement at the joint. With time and use, joints wear down. In osteoarthritis, a group of common joint disorders, cartilage repair does not keep up with degeneration, and the articular cartilage wears out. Inflammation of the joint capsule may occur, and bone outgrowths, or spurs, may develop. In rheumatoid arthritis, an autoimmune disease, the synovial membrane thickens and becomes inflamed. Synovial fluid accumulates, causing pressure, pain, stiffness, and progressive deformity, leading to loss of function.

Relate the structure of insect flight muscles to their function. Describe the structure of skeletal muscles and their antagonistic actions. List, in sequence, the events that take place during muscle contraction. Compare the roles of glycogen, creatine phosphate, and ATP in providing energy for muscle contraction. 11 Compare the structures and functions of the three types of skeletal muscle fibers. 12 Describe factors that influence contraction.

Most animals have the ability to move from place to place. Throughout most of the animal kingdom, a muscular system generates the mechanical forces and motion necessary for locomotion and for manipulation of objects. Muscle also powers many physiological actions necessary to maintain homeostasis. Many animals have digestive systems that push food along with peristaltic contractions and hearts that pump internal circulating fluids. Some animals also have blood vessels that maintain their pressure with gentle squeezing. Animals with very simple body plans do not have muscle tissue, but all eukaryotic cells do contain the contractile protein actin. The major component of microfilaments (see Fig. 4-28), actin is important in many cell processes, including amoeboid movement and attachment of cells to surfaces. In most cells, actin is functionally associated with the contractile protein myosin. Actin and myosin are most highly organized in muscle fibers.

Invertebrate muscle varies among groups Earlier in the chapter, we discussed the contractile cells of the hydrostatic skeleton of Hydra and other cnidarians. In most other animal groups, muscle is a specialized tissue organized into definite layers, or straplike bands. Some invertebrate phyla have skeletal and smooth muscle. Bivalve mollusks, such as clams, have two sets of muscles for opening and closing the shell. Their specialized smooth muscle, which is capable of slow, sustained contraction at low energy cost, keeps the two shells tightly closed for long periods, even weeks at a time. Striated muscle, which contracts rapidly, is used to swim and to shut the shell quickly when the mollusk is threatened. Arthropod muscles are striated, even in the walls of the digestive tract.

Insect flight muscles are adapted for rapid contraction Insects were the first animals in which flight evolved, an adaptation that has contributed to their impressive biological success

(see Chapter 31). Just how insects fly has been an aerodynamic Vertebrate skeletal muscles act mystery, and scientists are working to understand their remarkable antagonistically to one another ability to maneuver. Insect flight muscles contract more rapidly Skeletal muscles produce movements by pulling on tendons, than any other known muscle, up to 1000 contractions per second! tough cords of connective tissue that anchor muscles to bone. Not surprisingly, insect flight muscles in action have the highest Tendons then pull on bones. Skeletal muscles, or their tendons, known metabolic rate of any muscle tissue. pass across a joint and are attached to the bones on each side of the As you might imagine, insect flight muscles are structurally joint. When the muscle contracts, it pulls one bone toward or away well adapted to their function. They contain more mitochondria from the bone with which it articulates. than any other known variety of muscle. Their mitochondria also Because muscles can only contract, they can only pull; they have more cristae (folds of their inner membranes) and can concannot push. Muscles act antagonistically to one another, which sume oxygen twice as fast as mammalian mitochondria. Insect means that the movement produced by one can be reversed by anmuscles are also elaborately infiltrated with tracheae, tiny air-filled other. Th e biceps muscle, for example, flexes (bends) your arm, tubes that carry oxygen directly to each fiber. whereas contraction of the triceps muscle extends it (FIG. 40-7). Many insects must warm up before they fly because an increase Th us, the biceps and triceps work antagonistically. in body temperature increases the rate of ATP synthesis. Butterflies Th e muscle that contracts to produce a particular action is and some other insects bask in the sun before they fly. Honeybees known as the agonist. Th e muscle that produces the opposite “shiver” their muscles to increase body temperature. movement is the antagonist. When the agonist is contracting, the In the common blowfly, the wings may beat at 120 cycles per antagonist is relaxed. Generally, movements are accomplished by second. Yet in the same blowfly, the neurons that innervate those fugroups of muscles working together, so several agonists and sevriously contracting flight muscles are delivering impulses to them eral antagonists may take part in any action. Note that muscles that at the astonishingly low frequency of three per second. In fact, are agonists in one movement may serve as antagonists in another. about 75% of flying insect species beat their wings too rapidly for Some of the superfi cial skeletal muscles of the human body are each contraction to be controlled by a signal from a motor neuron. FIGURE 40-8 . shown in Instead, they have asynchronous, or indirect, muscle contractions in which muscle contraction is not synchronized with signals from motor neurons. In insects with asynchronous muscle contractions, the striated flight muscles do not attach directly to the wings. They attach to the flexible portions of the exoskeleton that articulate with the wings. The mechanical properties of this musculoskeletal arrangement provide the stimuli for contraction by stretching the muscle fibers at a Biceps relaxes high frequency. Each contraction of the muscles “dimples” the exoskeleton in association with a downstroke, and sometimes, depending on the Triceps contracts exact arrangement of the muscles, on the upstroke as well. When the dimple springs back into resting position, the muscles attached to it stretch. Triceps relaxes The stretching immediately initiates another contraction, and the cycle repeats. Thus, the deformation of the exoskeleton is transmitted as a force to Biceps contracts the wings, which beat so fast that we may perceive the sounds as a musical tone. However, nerve impulses are needed to maintain contraction. Somehow insects create lift that is 20 times or more their body weight. A central question has been how their flapping wings generate enough force to keep them airborne. Insect flight involves much more than just flapping the wings up and down. The flapping motion of the insect wing (a) Flexion. When the biceps muscle (b) Extension. When the triceps changes direction and speed, and upstrokes altercontracts, the arm flexes (bends). muscle contracts, the arm extends. nate with downstrokes at very high rates. At each shift of stroke, the wing rotates about its long axis FIGURE 40-7 Animated Muscle action and tilts to just the right angle for the new direcThe biceps and triceps muscles function antagonistically. tion of motion.

Muscles that flex fingers

Facial muscles Platysma

Sternocleidomastoid Trapezius

Latissimus dorsi Rectus abdominis

Clavicle Deltoid Pectoralis major Biceps brachii

Linea alba External oblique

Brachialis

Gluteus medius

Wrist and finger flexors

Gracilis Sartorius Quadriceps femoris

Sternocleidomastoid

Biceps brachii

Trapezius Deltoid

Brachialis

Triceps brachii

Latissimus dorsi

Brachioradialis

External oblique

Muscles that flex fingers

Gluteus maximus Hamstring muscles

Triceps brachii

Gracilis Semitendinosus

Patella

Biceps femoris Semimembranosus

Gastrocnemius Tibialis anterior Soleus

Gastrocnemius

Tibia

Soleus Achilles tendon Calcaneus

(a) Anterior view

(b) Posterior view

FIGURE 40-8 Some superficial muscles of the human body The labels in boldface are bones.

A vertebrate muscle may consist of thousands of muscle fibers Skeletal muscle is the most abundant tissue in the vertebrate body. Its elongated cells, called muscle fibers, are organized in bundles wrapped by connective tissue. The biceps in your arm, for example, consists of thousands of individual muscle fibers and their connective tissue coverings. Each striated muscle fiber is a long, cylindrical cell with many nuclei (FIG. 40-9). The plasma membrane, known as the sarcolemma in a muscle fiber, has multiple inward extensions that form a set of T tubules (transverse tubules). The cytoplasm of a muscle fiber is called sarcoplasm, and the endoplasmic reticulum is called the sarcoplasmic reticulum.

Myofibrils, which are threadlike structures, run lengthwise through the muscle fiber. They consist of even smaller structures, the myofilaments, or simply filaments. There are two types of myofilaments: myosin and actin filaments. Myosin filaments are thick, consisting mainly of the protein myosin. The thin actin filaments consist mostly of the protein actin; they also contain the proteins tropomyosin and troponin, which regulate the actin filament’s interaction with myosin filaments. Myosin and actin filaments are organized into repeating units called sarcomeres, the basic units of muscle contraction. Hundreds of sarcomeres connected end to end make up a myofibril. Sarcomeres are joined at their ends by an interweaving of filaments called the Z line. Each sarcomere consists of overlapping myosin and actin filaments. The filaments overlap lengthwise in the muscle

KEY POINT

A skeletal muscle fiber contains threadlike myofibrils that consist of actin and myosin filaments; the filaments are organized into repeating units called sarcomeres.

Muscle fibers

Biceps muscle

(a) A muscle such as the biceps in the arm consists of many fascicles (bundles) of muscle fibers.

(b) A fascicle wrapped in a connective tissue covering.

Sarcolemma

Sarcoplasmic reticulum

(c) Part of a muscle fiber showing the structure of myofibrils. The Z lines mark the ends of the sarcomeres.

T tubule Mitochondria Nucleus

Myofibril

Z line

Myofilaments

(d) TEM of striated muscle.

25 μm

(e) LM showing striations.

FIGURE 40-9 Animated Muscle structure The regular pattern of overlapping filaments gives skeletal and cardiac muscle their striated appearance.

Ed Reschke/Peter Arnold

1 μm

Bloom and Fawcett Textbook of Histology

Sarcomere

fibers, producing the pattern of transverse bands or striations characteristic of striated muscle (FIG. 40-10; also see Fig. 40-9). The bands are designated by the letters A, H, and I. The I band consists of parts of actin filaments of two adjacent sarcomeres. The A band is the wide, dark region that includes overlapping myosin and actin filaments. Within the A band is a narrow, light area, the H zone, made up exclusively of myosin filaments; the actin filaments do not extend into this region.

Contraction occurs when actin and myosin filaments slide past one another Muscle contraction occurs when the sarcomeres, and thus the muscle fibers, shorten. This explanation of muscle contraction, known as the sliding filament model, was developed in the 1950s by two British biologists, Hugh Huxley and Andrew Huxley. (The Huxleys are not related. They worked separately, and independently came to the same general conclusions at about the same time.) Thanks to the Huxleys, we now understand that the muscle shortens as the actin and myosin filaments slide past one another, increasing their overlap. You might think of an extension ladder. The overall ladder length changes as the ends get closer or farther apart, but the length of each ladder section stays the same. The I band and H zone decrease in length, but neither the actin nor myosin filaments themselves shorten. Motor neurons transmit messages from the brain or spinal cord to muscle fibers. When a motor neuron transmits a message, it releases the neurotransmitter acetylcholine into the synaptic cleft, a small space between the motor neuron and each muscle fiber. Acetylcholine binds with receptors on each muscle fiber, causing depolarization, a change in the distribution of electric charge across its sarcolemma. Depolarization can cause an electrical signal, or action potential, to be generated in the muscle fiber. In a muscle fiber, an action potential is a wave of depolarization that travels along the sarcolemma and into the system of T-tubule membranes. Depolarization of the T tubules opens calcium channels in the sarcoplasmic reticulum, and stored calcium ions are released into the myofibrils. Calcium ions bind to the protein troponin on the actin filaments, which changes the shape of the troponin. This change results in the troponin pushing the tropomyosin away from the active sites on the actin filament (FIG. 40-11). These active sites, also called myosin-binding sites, are now exposed. One end of each myosin molecule is folded into two globular structures called heads. The rounded heads of the myosin molecules extend away from the body of the myosin filament. Each myosin molecule also has a long tail that joins other myosin tails to form the body of the thick filament. ATP is bound to the myosin when the muscle fiber is at rest (not contracting). Myosin is an adenosine triphosphatase (ATPase), an enzyme that splits ATP to form ADP and inorganic phosphate (Pi). Myosin converts the chemical energy of ATP into the mechanical energy of sliding filaments. According to the current model of muscle contraction, when ATP is split, the ADP and Pi initially remain attached to the myosin head. The myosin head (with ADP and Pi still bound to it) is in an energized state; it is “cocked.” The myosin head binds to an exposed active site on the actin filament, forming a cross bridge

linking the myosin and actin filaments. The inorganic phosphate is then released, which triggers a conformational change in the myosin head. The myosin head bends about 45 degrees, in a flexing motion. This movement, the power stroke, pulls the actin filament closer to the center of the sarcomere. During the power stroke, the ADP is released. A new ATP must bind to the myosin head before the myosin can detach from the actin. If sufficient calcium ions are present, the cycle begins anew. Energized once again, myosin heads contact a second set of active sites on the actin filament; this next set is farther down the filament, closer to the end of the sarcomere. The process repeats with a third set of active sites, and so on. This series of stepping motions pulls the actin filaments toward the center of the sarcomere. Each time the myosin heads attach, move 45 degrees, detach, and then re-attach farther along the actin filament, the muscle shortens. One way to visualize this process is to imagine the myosin heads engaging “hand over hand” with the actin filaments. When many sarcomeres contract simultaneously, they contract the muscle as a whole. The sequence of events in muscle contraction can be summarized as follows: 1. A motor neuron releases acetylcholine, which binds with receptors on the muscle fiber; binding causes depolarization and generation of an action potential. 2. The action potential spreads through T tubules, triggering Ca2+ release from the sarcoplasmic reticulum. 3. Ca2+ binds to troponin, which then undergoes a conformational change; active sites on actin filaments are exposed. 4. ATP (attached to myosin) is split, and the energized myosin head is cocked; it binds to the active site on the actin filament, forming a cross bridge. 5. The release of Pi from the myosin head triggers the power stroke. 6. During the power stroke, myosin heads pull actin filament toward the center of the sarcomere, shortening the muscle; ADP is released. 7. The myosin head binds ATP and detaches from actin. If the concentration of Ca2+ is high enough, the sequence repeats from step 3. When impulses from the motor neuron cease, the enzyme acetylcholinesterase in the synaptic cleft inactivates acetylcholine. Muscle fibers return to their resting state. Calcium ions are pumped back into the sarcoplasmic reticulum by active transport, a process that requires ATP. Calcium removal is the key step in muscle relaxation. Without Ca2+, tropomyosin once again covers active sites on the actin filaments. The actin filaments slide back to their original position, and the muscle relaxes. This entire series of events happens in milliseconds.

ATP powers muscle contraction Muscle cells often perform strenuously and need large amounts of energy. As noted, the immediate source of energy for muscle contraction is ATP. Energy stored in ATP molecules powers cocking, attachment of the myosin heads to the actin filaments, flexion, and release of the myosin heads. Note that energy is needed not only

KEY POINT

A muscle contracts when actin and myosin filaments slide past one another. The length of each filament remains the same.

Cross bridges Actin (thin filament) Myosin (thick filament)

(a) Cross section of a myofibril. Note the arrangement of actin and myosin filaments. Each myosin filament is surrounded at each end by six actin filaments.

Sarcomere A band

(c) Muscle contraction. The I band and H zone decrease in length as the filaments slide past one another. 1

H zone

(b) Relationship of a muscle fiber, myofibrils, and filaments. Part of a muscle fiber showing the location of the filaments in a myofibril. I band

Z line

Muscle fiber in relaxed state. Note that I band and H zone are relatively wide. Actin (thin) filament Myosin (thick) filament

H. E. Huxley

Cross bridges

Muscle fiber contracting. Actin and myosin filaments slide past one another, increasing amount of overlap between actin and myosin filaments. Note that I band and H zone decrease in length. Filaments themselves do not become shorter.

3

Contracted muscle fiber. Actin filaments overlap, eliminating H zone. I bands also disappear as Z lines are pulled close to myosin filaments.

H. E. Huxley

2

Sarcomere

FIGURE 40-10 Animated The sliding filament model of muscle contraction Myosin and actin filaments are organized into repeating units called sarcomeres. As the sarcomeres become shorter, the muscle fiber short-

ens, but the size of the filaments does not change. (The I band consists of actin filaments of two adjacent sarcomeres.)

KEY POINT

The energy of ATP powers muscle contraction. During the power stroke, myosin heads move actin filaments toward the center of the sarcomere, which shortens the muscle.

1

Acetylcholine (released by motor neuron) combines with receptors on muscle fiber, causing depolarization and an action potential.

2

Neural impulse spreads through T tubules, stimulating Ca2+ release from sarcoplasmic reticulum.

ATP binding site Myosin filament ATP

2+

Ca

2+

Tropomyosin

Ca

Actin filament

Ca

Troponin

2+

P ADP

Ca

Ca2+

2+

2+

Ca

Ca2+

Active sites 4

3 Ca2+

bind to troponin, causing change in shape. Troponin pushes tropomyosin away, exposing active sites on actin filaments.

ATP is split. Myosin head, now cocked, binds to exposed active site, forming cross bridge.

If Ca2+ available

Ca2+

ATP

Ca2+

ADP 2+

Pi is released.

Actin-myosin complex binds ATP, and myosin detaches from actin.

ADP

6

P Ca

5 7

2+

Ca

Cross bridge flexes, and actin filament is pulled toward center of sarcomere. This movement is the power stroke. ADP is released.

FIGURE 40-11 Animated Muscle contraction: model of actin and myosin interactions Contraction results when actin filaments slide toward the center of individual sarcomeres of a myofibril. After step 7, the cycle repeats from step 3 if calcium ions are available. In the absence of calcium ions, the muscle relaxes.

for the pull exerted by the cross bridges but also for their TABLE 40-1 Characteristics of Skeletal Muscle Fibers release from each active site. Slow-Oxidative Fast-Glycolytic Fast-Oxidative Rigor mortis, the temporary but very marked muscle Characteristics Fibers Fibers Fibers stiffening that occurs after death, results from ATP depletion. When death occurs, cellular respiration ceases and Contraction speed Slow Fast Fast ATP is used up. However, at the time of death, many Rate of fatigue Slow Fast Intermediate muscle fibers are in the process of contraction, and cross Major pathway for Aerobic Glycolysis Aerobic ATP synthesis respiration respiration bridges have formed. Because ATP is essential for release Mitochondria Many Few Many of the myosin heads, cross bridges remain intact. Rigor Intensity of Low High Intermediate mortis does not persist indefinitely because the entire contraction contractile apparatus of the muscles eventually decomMyoglobin content High Low High poses, restoring pliability. The phenomenon is temperaColor of fi ber Red White Red ture dependent, so given the prevailing temperature, a medical examiner can estimate the time of death of a cadaver from its degree of rigor mortis. ATP molecules can provide energy for only a few Fast-glycolytic fibers generate a great deal of power and carry seconds of strenuous activity. Fortunately, muscle fibers have a out rapid movements but can sustain that activity for only a short backup energy storage compound, creatine phosphate, that can time. They are important in activities such as sprinting and weight be stockpiled. The energy stored in creatine phosphate is translifting. Fast-glycolytic fibers contract rapidly, have few mitochonferred to ATP as needed. But during vigorous exercise the supply dria, and obtain most of their energy from glycolysis. When their of creatine phosphate is quickly depleted. Muscle cells must reglycogen supply is depleted, they fatigue rapidly. These fibers have plenish their supplies of these energy-rich compounds. a low myoglobin content and are white. (They are sometimes reMuscle fibers store chemical energy in glycogen, a large polyferred to as white fibers.) People who are sedentary have more fastsaccharide formed from hundreds of glucose molecules. Glycogen glycolytic fibers than do physically fit individuals. With physical can be degraded, yielding glucose, which is then degraded in cellutraining, these fibers change to fast-oxidative fibers. lar respiration. When sufficient oxygen is available, enough energy Fast-oxidative fibers, also known as intermediate fibers, conis captured from glucose to produce needed quantities of ATP and tract rapidly and have an intermediate rate of fatigue. Like slowcreatine phosphate. oxidative fibers, they are rich in mitochondria and obtain most of During a burst of strenuous exercise, the circulatory system their ATP from aerobic respiration. They have a high myoglobin cannot deliver enough oxygen to keep up with the demand of the content and a red color. rapidly metabolizing muscle fibers. This results in an oxygen debt. Entire muscles may be specialized for quick or slow responses. Under these conditions, muscle fibers break down fuel molecules In chickens, for instance, the white breast muscles are efficient for anaerobically (without oxygen) for short periods. Lactic acid ferquick responses, perhaps because a short flight is an escape mechmentation is a method of generating ATP anaerobically, but not anism for chickens. However, they walk about on the ground all in great quantity (see Fig. 8-14). ATP depletion results in weaker day; the dark (red) meat of the leg and thigh is muscle specialized contractions and muscle fatigue. Accumulation of the waste prodfor more sustained activity. Birds that fly have red breast muscles uct lactic acid also contributes to muscle fatigue. Well-conditioned specialized to support sustained activity. athletes develop the ability to tolerate the high levels of lactic acid In humans, most muscles have a mixture of different types of generated during high-performance activity. The period of rapid fibers. The proportions of the types of fibers vary among individbreathing that generally follows strenuous exercise pays back the uals and from muscle to muscle in the same person. Aerobic exoxygen debt by consuming lactic acid. ercise, such as jogging or bicycling, increases endurance. Aerobic training increases capillary density, the number of mitochondria, The type of muscle fibers determines and the myoglobin content of muscle fibers. In strength trainstrength and endurance ing, for example, weight lifting, specific muscles are repeatedly Three major types of skeletal muscle fibers have been identified in contracted under heavy loads. This action increases the number vertebrates: slow-oxidative, fast-glycolytic, and fast-oxidative (TAof filaments in the muscle fibers, which increases the size of the BLE 40-1). Slow-oxidative fibers are well-adapted for endurance acmuscle. tivities such as swimming and running long distances, and for maintaining posture. They contract slowly and fatigue slowly. These fibers Several factors influence the strength require a steady supply of oxygen. They derive most of their energy of muscle contraction from aerobic respiration and are rich in mitochondria and capillaries. In addition to the types of muscle fibers making up the muscle, Slow-oxidative fibers are rich in myoglobin, a red pigment similar contraction of a whole muscle depends on the number of muscle to hemoglobin, which stores oxygen in red blood cells. Myoglobin fibers contracting; the tension developed by each fiber; frequency enhances rapid diffusion of oxygen from blood into muscles durof stimulation; the condition of the muscle, for example, whether ing strenuous muscle exertion. Because of their myoglobin, slowor not it is fatigued; and the load on the muscle. The more muscle oxidative fibers are red and are sometimes called red fibers.

fibers that contract, the greater the tension in the muscle. In a motor unit, a motor neuron is functionally connected with an average of about 150 muscle fibers (FIG. 40-12). However, some motor units have less than a dozen motor neurons, and others have several hundred. Each junction of a motor neuron with a muscle fiber is called a neuromuscular junction. Messages from the brain or

Cross section of spinal cord

Neuromuscular junctions

(a) The motor unit illustrated here shows only a single motor neuron fiber.

spinal cord activate motor units. The more motor units that are recruited, the stronger are the contractions. An important factor affecting the tension of contracting muscle fibers is the frequency of stimulation. In response to a single, brief electrical stimulus, skeletal muscle contracts with a single, quick contraction called a simple twitch. When a second stimulus is received before the first contraction is complete, the two twitches may add together, a process called summation (FIG. 40-13). Typically, skeletal muscle receives a series of separate stimuli timed very closely together. Summation results in a smooth, sustained contraction called tetanus. Depending on the identity and number of our muscle fibers tetanically contracting, we might thread a needle, rock a baby, or run a mile. When you lift a heavy object, you place a load on the muscles affected. As the weight of the load increases, the muscles contract more strongly (within limits). Even when you are not moving, your muscles are in a state of partial contraction known as muscle tone. At any given moment, some muscle fibers are contracted, stimulated by messages from motor neurons. Muscle tone is an unconscious process that keeps muscles prepared for action. When the motor nerve to a muscle is cut, the muscle becomes limp (completely relaxed, or flaccid) and eventually atrophies (decreases in size).

Smooth muscle and cardiac muscle are involuntary

Spinal nerve

Muscle

Don Fawcett/Science Source/Photo Researchers, Inc.

Motor nerve fiber Neuromuscular junction

Part of muscle fiber

10 μm

(b) SEM of some of the fibers in a motor unit. Note how neurons branch to innervate all muscle fibers in the motor unit.

FIGURE 40-12 Animated A motor unit A motor unit consists of a motor neuron and the muscle fibers that it innervates.

The three types of vertebrate muscle—skeletal, smooth, and cardiac—are compared in Table 39-3. All three types of muscle have thin actin filaments that slide along thick myosin filaments in response to an increase in calcium ion concentration. All three muscle types use ATP to power contraction. However, each type of muscle is specialized for particular types of responses. Neither smooth muscle nor cardiac muscle is under voluntary control. Smooth muscle is not attached to bones but instead forms tubes that squeeze like the muscle tissue in the body wall of the earthworm. Smooth muscle often contracts in response to simple stretching, and its contraction tends to be sustained. It is well adapted to performing such tasks as regulating blood pressure by sustained contraction of the walls of the arterioles. Although smooth muscle contracts slowly, it shortens much more than striated muscle does; it squeezes impressively. Smooth muscle is not striated because its actin and myosin filaments are not organized into myofibrils or into sarcomeres. The fibers of smooth muscle tissue function as a unit because they are connected by gap junctions (see Chapter 5). Gap junctions permit electrical signals to pass rapidly from fiber to fiber. Although smooth muscle contraction is basically similar to skeletal muscle contraction (occurring by a sliding filament mechanism), the cross bridges in smooth muscle remain in the attached state longer. For this reason, less ATP is required to maintain a high level of force. Cardiac muscle contracts and relaxes in alternating rhythm, propelling blood with each contraction. Sustained contraction of cardiac muscle would be disastrous! Like smooth muscle fibers, cardiac muscle fibers are electrically coupled by gap junctions

Tetanus

Tension

3

2

Single twitch

Stimulation ends

Twitch summation

1

0 Time

(a) When a muscle fiber is allowed to completely relax and is then stimulated a second time, the second twitch is the same magnitude as the first twitch.

(b) When a muscle fiber is stimulated a second time before it has completely relaxed, the second twitch is added on to the first twitch. The result is summation.

(c) A volley of rapid stimuli does not allow the muscle fiber to relax between stimuli. A sustained contraction, called tetanus, occurs.

FIGURE 40-13 Animated Summation and tetanus

(called intercalated discs). Each heartbeat is initiated by a pacemaker, a mass of specialized cardiac muscle. A conduction system transmits the signal throughout the cardiac muscle, and the heart beats in a regular rhythm independent of a nerve supply. However, the heart rate is regulated, according to the needs of the body, by neurons in the cardiac centers in the medulla of the brain (discussed in Chapter 44).

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Review Describe a skeletal muscle fiber. What are its two types of filaments? What events occur when a muscle fiber contracts (begin with release of acetylcholine and include cross-bridge action)? What is the role of ATP in muscle contraction? What are the functions of creatine phosphate and glycogen? What is summation?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

40.1 (page 843) 1 Compare the functions of the epithelial coverings of invertebrates and vertebrates. ■ Epithelial tissue protects underlying tissues and may be specialized for sensory or respiratory functions. The outer epithelium may be specialized to secrete antibacterial or poisonous substances. In many invertebrates, the outer epithelium is specialized to ■ secrete a protective cuticle, or shell. ■ The integumentary system of vertebrates includes the skin and structures that develop from it. Mammalian skin includes hair, claws or nails, sweat glands, oil glands, and sensory receptors. 2 Relate the structure of vertebrate skin to its functions. ■ The feathers of birds and the hair of mammals form an insulating layer that helps maintain a constant body temperature. Mammalian skin protects the body from the wear and tear that occurs as it interacts with the outer environment. Stratum corneum, the most superficial layer of the epidermis, consists of dead cells, filled with keratin, a protein that gives mechanical strength to the skin and reduces water loss. Cells in stratum basale of the epidermis divide and are pushed upward

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Epidermis

toward the skin surface. These cells mature, produce keratin, and eventually die and slough off. The dermis consists of dense, fibrous connective tissue. In birds and mammals, the dermis rests on a layer of subcutaneous tissue composed largely of insulating fat.

Stratum corneum Stratum basale Dermis

Subcutaneous tissue

40.2 (page 844) 3 Compare the structure and functions of different types of skeletal systems, including the hydrostatic skeleton, exoskeleton, and endoskeleton. ■ The skeletal system supports and protects the body and transmits mechanical forces generated by muscles. Many soft-bodied invertebrates have a hydrostatic skeleton; fluid in a closed body compartment is used to transmit forces generated by contractile cells or muscle. ■ Exoskeletons are characteristic of mollusks and arthropods. The arthropod skeleton, composed partly of chitin, is jointed for flexibility and adapted for many lifestyles. This nonliving skeleton does not grow, making it necessary for arthropods to undergo ecdysis (molt) periodically. ■ The endoskeletons of echinoderms and chordates consist of living tissue and therefore can grow. 4 Describe the main divisions of the vertebrate skeleton and the bones that make up each division. ■ The vertebrate skeleton consists of an axial skeleton and an appendicular skeleton. The axial skeleton consists of skull, vertebral column, rib cage, and sternum. The appendicular skeleton consists of bones of the limbs, pectoral girdle, and pelvic girdle. 5 Describe the structure of a typical long bone and differentiate between endochondral and intramembranous bone development. ■ A typical long bone consists of a thin outer shell of compact bone surrounding the inner spongy bone and a central cavity that contains bone marrow. ■ Long bones develop from cartilage templates during endochondral bone development. Other bones, such as the flat bones of the skull, develop from a noncartilage connective tissue model by intramembranous bone development. ■ Osteoblasts, cells that produce bone, and osteoclasts, cells that break down bone, work together to shape and remodel bone. Learn more about the human skeletal system and a typical long bone by clicking on the figures in CengageNOW.

6 Compare the main types of vertebrate joints. ■

Joints are junctions of two or more bones. Ligaments are connective tissue bands that connect bones and limit movement at the joint. The sutures of the skull are immovable joints. Joints between vertebrae are slightly movable joints. A freely movable joint is enclosed by a joint capsule lined with a membrane that secretes synovial fluid.

40.3 (page 848) 7 Relate the structure of insect flight muscles to their function. Large numbers of mitochondria and tracheae (air tubes) present in insect flight muscles support the high metabolic rate required for flight. 8 Describe the structure of skeletal muscles and their antagonistic actions. A muscular system is found in most invertebrate phyla and in ■ vertebrates. As muscle tissue contracts (shortens), it moves body parts by pulling on them. ■ Vertebrate skeletal muscles pull on tendons, connective tissue cords that attach muscles to bones. When a muscle contracts, it pulls a bone toward or away from the bone with which it articulates. Skeletal muscles act antagonistically to one another. The muscle ■ that produces a particular action is the agonist; the antagonist produces the opposite movement. ■ A skeletal muscle such as the biceps is an organ made up of hundreds of muscle fibers. Each fiber consists of threadlike myofibrils composed of smaller myofilaments, or simply filaments. The striations of skeletal muscle fibers reflect the overlapping of their actin

filaments and myosin filaments. A sarcomere is a contractile unit of actin (thin) and myosin (thick) filaments. 9 List, in sequence, the events that take place during muscle contraction. ■ Acetylcholine released by a motor neuron binds with receptors on the surface of a muscle fiber. This may cause depolarization of the sarcolemma and transmission of an action potential. The action potential spreads through the T tubules, releasing calcium ions from the sarcoplasmic reticulum. ■ Calcium ions bind to troponin in the actin filaments, causing the troponin to change shape. Troponin pushes tropomyosin away from the binding sites on the actin filaments. ■ ATP binds to myosin; ATP is split, putting the myosin head in a highenergy state (it is “cocked”). Energized myosin heads attach to the exposed binding sites on the actin filaments, forming cross bridges that link the myosin and actin filaments. ■ After myosin attaches to the actin filament, the cross bridge flexes as phosphate is released. This is the power stroke, which pulls the actin filament toward the center of the sarcomere. ADP is released during the power stroke. ■ The myosin head binds a new ATP, which lets the myosin head detach from the actin. As long as the calcium ion concentration remains elevated, the new ATP is split and the sequence repeats. The myosin reattaches to new active sites so that the filaments are pulled past one another, and the muscle continues to shorten. 10 Compare the roles of glycogen, creatine phosphate, and ATP in providing energy for muscle contraction. ■ ATP is the immediate source of energy for muscle contraction. ATP hydrolysis provides the energy to “cock” the myosin. Muscle tissue has an intermediate energy storage compound, creatine phosphate. Glycogen is the fuel stored in muscle fibers. 11 Compare the structures and functions of the three types of skeletal muscle fibers. ■ Slow-oxidative fibers are specialized for endurance activities. They contract slowly, fatigue slowly, are rich in mitochondria, and obtain most of their ATP from aerobic respiration. These fibers have a red color because they have a high content of myoglobin, a red pigment that stores oxygen. ■ Fast-glycolytic fibers generate a great deal of power for a brief period. These white fibers contract rapidly, fatigue quickly, have few mitochondria, and use glycolysis as a major pathway for ATP synthesis. ■ Fast-oxidative fibers are specialized for rapid response. They contract fast, have an intermediate rate of fatigue, are rich in mitochondria, and obtain most of their ATP from aerobic respiration. They have a high myoglobin content and a red color.

12 Describe factors that influence contraction. ■

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Contraction of a whole muscle depends on several factors, including the number of muscle fibers contracting, tension developed by each fiber, frequency of stimulation, condition of the muscle, and load on the muscle. A motor unit consists of all of the skeletal muscle fibers stimulated by a single motor neuron. Messages from the brain or spinal cord activate motor units. The more motor units that are recruited, the stronger are the contractions. When activated by a brief electrical stimulus, skeletal muscle responds with a simple twitch. When a second stimulus is received before the first contraction is complete, the two twitches can add together, a process called summation. Typically, skeletal muscle is stimulated by a series of separate stimuli timed close together and responds with a smooth, sustained contraction called tetanus. Muscle tone is the state of partial contraction characteristic of muscles. Explore muscle action, structure, and contraction as well as summation and tetanus by clicking on the figures in CengageNOW.

T E S T YO U R U N D E R S TA N D I N G 1. In vertebrate skin, cells actively divide in (a) stratum basale (b) stratum corneum (c) the dermis (d) the layer with cells that contain keratin (e) stratum basale and stratum corneum 2. An endoskeleton (a) is typically composed of dead tissue (b) is characterized by fluid in a closed compartment (c) is found mainly in echinoderms and vertebrates (d) is typical of echinoderms and arthropods (e) requires the animal to molt 3. Which of the following is not part of the vertebrate axial skeleton? (a) skull (b) vertebral column (c) pelvic girdle (d) rib cage (e) sternum 4. The thin outer shell of a long bone is made of (a) compact bone (b) spongy bone (c) epiphyses (d) cartilage (e) mainly chondrocytes 5. In endochondral bone development, (a) osteoclasts produce bone (b) joints connect fibers (c) the skeleton consists of cartilage (d) bones develop from cartilage templates (e) bones form in noncartilage connective tissue 6. All animals have (a) muscles (b) actin (c) bones (d) endoskeleton or exoskeleton (e) dermis

8. Myosin binds to actin, forming a cross bridge. What happens next? (a) acetylcholine is released (b) calcium ions stimulate exposure of active sites (c) filaments slide past one another, and the muscle fiber shortens (d) myosin is activated (e) Pi is released, and the cross bridge flexes 9. Calcium ions are released from the sarcoplasmic reticulum. What happens next? (a) acetylcholine is released (b) active sites on the actin filaments are exposed (c) filaments slide past one another, and the muscle fiber shortens (d) myosin is activated (e) Pi is released, and the cross bridge flexes 10. When skeletal muscle is stimulated by a series of closely timed separate stimuli, (a) white fibers respond (b) a simple twitch occurs (c) it responds with a smooth, sustained contraction called tetanus (d) red fibers respond (e) summation is inhibited 11. Insect flight muscles (a) work best at low temperature (b) evolved after bird wings (c) do not create much lift (d) have a very high metabolic rate (e) are typically attached directly to the wing and make up the bulk of the wing 12. Label the diagram. (Refer to Fig.40-9c to check your answers.)

7. An energy storage compound that can be stockpiled in muscle cells for short-term use is (a) myoglobin (b) ADP (c) troponin (d) myosin (e) creatine phosphate

CRITICAL THINKING 1. What are some examples of hydrostatic support in plants? 2. Why is it important that a muscle be able to shift functions, sometimes acting as an agonist and at other times acting as an antagonist? 3. What is the functional relationship between skeletal and muscle tissues? 4. EVOLUTION LINK. Very different skeletal systems have evolved in arthropods and vertebrates. What are some benefits and some

disadvantages of each type of skeleton? In what way is the insect skeleton an adaptation to the insect lifestyle? 5. ANALYZING DATA. Examine Figure 40-13. Based on the summation of two stimuli shown in this graph, draw a curve showing summation when the muscle fiber is stimulated three times. Additional questions are available in CengageNOW at www.cengage.com/ login.

41

Neural Signaling

Nancy Kedersha/UCLA/Science Photo Library/ Photo Researchers, Inc.

Specialized cells in the nervous system. This immunofluorescent LM of mammalian spinal cord tissue includes several astrocytes (green). The blue dots are the nuclei of other glial cells. The smaller cells (dark pink) are neurons; their cell bodies appear light pink.

KEY CONCEPTS

A

41.1 Neural signaling involves reception, transmission,

changes in its environment. Most animals have a nervous system that

integration, and action by effectors.

takes in information, transmits it to the spinal cord and brain where it is

41.2 Neurons are specialized to receive stimuli and trans-

integrated, and then responds. The nervous system is composed mainly

mit signals; glial cells support, protect, and nourish neurons and can modify neural signals.

of two specialized types of cells—neurons and glial cells (see photograph).

41.3 The resting potential of a neuron is maintained by differences in concentrations of specific ions inside the cell relative to the extracellular fluid and by selective permeability of the plasma membrane to these ions. Depolarization of the neuron plasma membrane to threshold level generates an action potential, an electrical signal that travels as a wave of depolarization along the axon.

41.4 Most neurons signal other cells by releasing neuro-

n animal’s ability to survive and to maintain homeostasis depends largely on how effectively it detects and responds to stimuli—

Just how animals respond to stimuli depends on how their neurons are organized and connected to one another. A single neuron in the vertebrate brain may be functionally connected to thousands of other neurons. The nervous system is the principal regulatory system in animals. Regulation requires communication, and the nervous system transmits information to and from all parts of the body. As you will learn, the endocrine system works with the nervous system to regulate many behaviors

transmitters at chemical synapses.

and physiological processes. In general, the endocrine system provides

41.5 During integration, incoming neural signals are

relatively slow and long-lasting regulation, whereas the nervous system

summed; temporal and spatial summation can bring a neuron to threshold level.

responds rapidly, but briefly.

41.6 Neurons are organized in specific pathways called

Active areas of research include neurotransmitters, the chemical messen-

neural circuits, which, in turn, are organized to form neural networks.

gers used by neurons to signal other neurons, and the receptors that bind

Neurobiology is one of the most exciting areas of biological research.

with the neurotransmitters. Some neurobiologists are focused on how neural networks in the brain function in cognitive and emotional experience. For example, researchers have recently identified an area of the

prefrontal cortex important in self-control and decision-making. They have also identified neural circuits that appear to be responsible for our experiences of physical pain and pleasure.

External stimulus (e.g., vibration, movement, light, odor)

Internal stimulus (e.g., change in blood pH or blood pressure)

A very active area of research focuses on neurogenesis, the production of new neurons. One of the most time-honored views about nervous system function was that the brain of mammals cannot generate new neurons after birth. Nobel laureate Santiago Ramon y Cajal promoted this concept at the beginning of the 20th century, and the idea that we are born

RECEPTION Detection by external sense organs

Detection by internal sense organs

with all the brain cells we will ever have was widely held for almost a century. Another enduring idea was that glial cells, the most common cells in the brain, simply provide support services for neurons. As we will discuss in this chapter, both of these concepts have been shown to be incorrect.

TRANSMISSION Sensory (afferent) neurons transmit information

In this chapter we focus on how information is communicated by neurons and glial cells. In Chapter 42 we discuss nervous systems, and in Chapter 43 we explore sensory reception.

Central Nervous System (brain and spinal cord) INTEGRATION

41.1 NEURAL SIGNALING: AN OVERVIEW ■ ■ LEARNING OBJECTIVE 1

Describe the processes involved in neural signaling: reception, transmission, integration, and action by effectors.

Interneurons sort and interpret information

TRANSMISSION Motor (efferent) neurons transmit impulses

Thousands of stimuli constantly bombard an animal. Survival depends on identifying these stimuli and responding effectively. Information about each stimulus must be transmitted to the nervous system and then information from the nervous system must be transmitted to the muscles (and glands). AniACTION BY EFFECTORS mals respond to changes in temperature, sound, light, odors, (muscles and glands) and movement that may indicate the presence of a predator or of prey. Internal signals, such as hunger, pain, fluctuations in blood pressure, or changes in blood sugar concentration, also e.g., respiration demand responses. In most animals, responses to stimuli dee.g., animal rate increases; runs away pend on neural signaling, information transfer by networks blood pressure of nerve cells, called neurons. In most animals, neural signalrises ing involves four processes: reception, transmission, integration, and action by effectors (muscles or glands). Reception, the process of detecting a stimulus, is the job FIGURE 41-1 Animated Response to a stimulus of neurons and of specialized sensory receptors such as those in Whether a stimulus originates in the outside world or inside the body, information the skin, eyes, and ears (FIG. 41-1). Transmission is the promust be received; transmitted to the central nervous system; integrated; and then cess of sending messages along a neuron, from one neuron to transmitted to effectors, muscles or glands that carry out some action, the actual response. another or from a neuron to a muscle or gland. In vertebrates, a neural message is transmitted from a receptor to the central nervous system (CNS), which consists of the brain and spinal cord. put. Integration involves sorting and interpreting incoming senNeurons that transmit information to the CNS are called afferent sory information and determining the appropriate response. (meaning “to carry toward”) neurons, or sensory neurons. Neural messages are transmitted from the CNS by efferent Afferent neurons generally transmit information to interneu(meaning “to carry away”) neurons to effectors—muscles and rons, or association neurons, in the CNS. Most neurons, perhaps glands. Efferent neurons that signal skeletal muscle are called motor 99%, are interneurons. Their function is to integrate input and out-

neurons. The action by effectors is the actual response to the stimulus. Sensory receptors and afferent and efferent neurons are part of the peripheral nervous system (PNS). In summary, information flows through the nervous system in the following sequence:

amine the structure of a common type of neuron, the multipolar neuron shown in FIGURE 41-2. The largest portion of the neuron, the cell body, contains the nucleus, the bulk of the cytoplasm, and most of the organelles. Typically, two types of processes project from the cell body of a multipolar neuron. Numerous dendrites extend from one end, and a long, single axon projects from the opposite end. Dendrites are typically short, highly branched processes specialized to receive stimuli and send signals to the cell body. The cell body integrates incoming signals. Although microscopic in diameter, an axon may be 1 m (more than 1 yard) or more in length and may divide, forming branches called axon collaterals. The axon conducts nerve impulses away from the cell body to another neuron or to a muscle or gland. At its end the axon divides, forming many terminal branches that end in synaptic terminals. The synaptic terminals release neurotransmitters, chemicals that transmit signals from one neuron to another or from a neuron to an effector. The junction between a synaptic terminal and another neuron (or effector) is called a synapse. Typically, a small space exists between these two cells. In vertebrates, the axons of many neurons outside the CNS are surrounded by a series of Schwann cells. The plasma membranes of these glial cells contain myelin, a white, fatty material. Schwann cells wrap their plasma membranes around the axon, forming an insulating covering called the myelin sheath. Gaps in the myelin sheath, called nodes of Ranvier, occur between successive Schwann cells. At these points the axon is not insulated with myelin. Axons more than 2 μm in diameter have myelin sheaths and are described as myelinated. Those of smaller diameter are generally unmyelinated.

reception by sensory receptor ¡ transmission by afferent neuron ¡ integration by interneurons in CNS ¡ transmission by efferent neuron ¡ action by effectors

Review ■

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Imagine you are swimming and you suddenly spot a shark fin moving in your direction. What processes must take place within your nervous system before you can make your escape? What happens after neural signals are integrated in the CNS?

41.2 NEURONS AND GLIAL CELLS ■ ■ LEARNING OBJECTIVES 2 3

Describe the structure of a typical neuron and give the function of each of its parts. Name the main types of glial cells and describe the functions of each.

Neurons and glial cells are unique to the nervous system. Neurons are specialized to receive and send information. Glial cells support and protect neurons and carry out many regulatory functions.

Neurons receive stimuli and transmit neural signals The neuron is highly specialized to receive and transmit information. Neurons produce and transmit electrical signals called nerve impulses, or action potentials. The neuron is distinguished from all other cells by its long processes (cytoplasmic extensions). Ex-

Certain regions of the CNS produce new neurons At the beginning of the chapter, we described Cajal’s idea that neurogenesis, the production of new neurons, does not occur in the CNS. The paradigm shift that has occurred with regard to

Dendrites covered with dendritic spines Synaptic terminals

Cytoplasm of Schwann cell Axon

Axon collateral

Cell body

Nucleus Myelin sheath

Nucleus Axon Nodes of Ranvier

Schwann cell

Terminal branches

FIGURE 41-2 Animated Structure of a multipolar neuron The cell body contains most of the organelles. Many dendrites and a single axon extend from the cell body. Schwann cells form a myelin sheath around the axon by wrapping their own plasma membranes around the neuron in jelly-roll fashion.

this view demonstrates the power and importance of the process of science. Cajal’s idea was so entrenched that for many years the researchers who attempted to study the issue were discouraged and discounted. Even images produced by an electron microscope showing neurons dividing were ridiculed and rejected. During the 1980s, studies on rhesus monkey brains appeared to support Cajal’s idea. No new neurons were found in the monkeys’ brains. The matter appeared settled. However, matters are almost never settled in science. Researchers investigating a variety of seemingly unrelated problems made observations that shed new light on the neurogenesis issue. For more than 25 years, Fernando Nottebohm at Rockefeller University has studied birdsong, including its neural basis. Nottebohm found that birds produce new neurons daily in the brain center that regulates their songs. His work was at first dismissed as a phenomenon peculiar to birds and not relevant to mammals. However, other investigators, most notably psychologist Elizabeth Gould at Princeton University, have provided additional data that together have falsified Cajal’s concept. Gould demonstrated neuron proliferation in primates. She and her research team also discovered that stress damages the brain and inhibits mitosis in neurons. They stressed newborn rats by separating them from their mothers for up to 3 hours a day. These rats did not learn how to cope with stress, and as adults, they had fewer new neurons in certain areas of their brains than control rats. Earlier investigations may have failed to show new neurons in the brain because the animals being studied were in a state of stress and neurogenesis was inhibited. Neurobiologists now agree that neurogenesis does occur regularly in the adult mammalian brain, at least in the olfactory bulb and hippocampus. The paradigm has shifted.

Axons aggregate to form nerves and tracts A nerve consists of hundreds or even thousands of axons wrapped together in connective tissue (FIG. 41-3). We can compare a nerve to a telephone cable. The individual axons correspond to the wires that run through the cable, and the sheaths and connective tissue coverings correspond to the insulation. Within the CNS, bundles of axons are called tracts or pathways rather than nerves. Outside the CNS, the cell bodies of neurons are usually grouped in masses called ganglia (sing., ganglion). Inside the CNS, collections of cell bodies are generally called nuclei rather than ganglia.

Glial cells play critical roles in neural function Glial cells collectively make up the neuroglia (literally, “nerve glue”). Until recently, glial cells were thought to be passive cells that simply support and protect the neurons. Armed with more sophisticated technology, researchers are discovering that glial cells communicate and carry out major regulatory functions. More than 75% of the cells in the human CNS are glial cells. They account for about 50% of the brain volume because they do not branch as extensively as neurons. Four types of glial cells are found in the vertebrate CNS: astrocytes, oligodendrocytes, ependymal cells, and microglia (TABLE 41-1). Astrocytes are star-shaped glial cells that provide physical support for neurons and provide neurons with nutrients. Astrocytes help regulate the composition of the extracellular fluid in the CNS by removing excess potassium ions. This action helps maintain normal neuron excitability. Some astrocytes position the ends of their long processes on blood vessels in the brain. In response,

Ganglion

Cell bodies

Artery

Vein

Axon 100 μm

(a) A nerve consists of bundles of axons held together by connective tissue. The cell bodies belonging to the axons of a nerve are grouped in a ganglion.

FIGURE 41-3 Animated Structure of a myelinated nerve

(b) SEM showing a cross section through a myelinated afferent nerve of a bullfrog.

E. R. Lewis / Biological Photo Service

Myelin sheath

TABLE 41-1

Functions of Glial Cells

Type of Glial Cell

Functions

Astrocytes

Physically support neurons Provide neurons with nutrients Remove excess K+, which helps regulate composition of extracellular fluid in CNS Induce blood vessels to form blood–brain barrier Communicate with one another and with neurons Induce synapse formation and strengthen activity of synapses Respond to neurotransmitters and help regulate neurotransmitter reuptake May be important in memory and learning Guide neurons during embryonic development

Capillary Astrocytes

Oligodendrocytes

Form myelin sheaths around neurons in CNS

Oligodendrocyte

Nerve fiber

Myelin sheath

Ependymal cells Cilia

Line cavities of CNS Help produce and circulate cerebrospinal fluid May function as neural stem cells

Ependymal cells

Microglia

Phagocytosis of bacteria and debris Release signaling molecules that mediate inflammation

endothelial cells lining the blood vessels form tight junctions (see Chapter 5) that prevent many substances in the blood from entering the brain tissue. This protective wall is the blood–brain barrier. Neurobiologists have shown that astrocytes are functionally connected throughout the brain. These interesting glial cells may participate in information signaling in the brain by coordinating activity among neurons. They communicate through gap junctions and with signaling molecules. Although astrocytes can generate only weak electrical signals, they communicate with one another and with neurons by means of chemical signals. Astrocytes appear to respond to neurotransmitters released by neurons. They also help regulate the reuptake of excess neurotransmitters from synapses. Neurobiologists have demonstrated that astrocytes induce synapse formation and can strengthen activity of synapses in the brain. Their actions appear to be important in learning and memory. Astrocytes play a role in development by guiding neurons to appropriate locations in the body. Neurobiologists have shown that certain astrocytes function as stem cells in the brain and spinal cord. These cells can give rise to new neurons, additional astrocytes, and certain other glial cells. Taken out of their normal environment in the adult mouse brain, astrocytes can give rise to cells of all germ layers (the embryonic tissue layers: ectoderm, mesoderm, and endoderm). Human astrocytes may someday be used to produce specific types of cells needed for treating various medical conditions. Oligodendrocytes are glial cells that envelop neurons in the CNS, forming insulating myelin sheaths around them. Because myelin is an excellent electrical insulator, its presence speeds transmission of neural impulses. In the neurological disease multiple sclerosis, which affects about 400,000 people in North America alone, patches of myelin deteriorate at irregular intervals along axons in the CNS and are replaced by scar tissue. This damage interferes with conduction of neural impulses, and the victim suffers loss of coordination, tremor, and partial or complete paralysis of parts of the body. Do not confuse oligodendrocytes with Schwann cells. Although sometimes considered glial cells, Schwann cells are located outside the CNS. Schwann cells form myelin sheaths around axons of many neurons that are part of the PNS. Ependymal cells are ciliated glial cells that line the internal cavities of the CNS. Ependymal cells help produce and circulate the cerebrospinal fluid that bathes the brain and spinal cord of vertebrates (discussed in Chapter 42). Investigators have recently suggested that ependymal cells might function as neural stem cells. Microglia are actually specialized macrophages (phagocytic cells that ingest and digest cell debris and bacteria). Microglia respond to signals from neurons and are important in mediating responses to injury or disease. These cells are found near blood vessels. When the brain is injured or infected, microglia multiply and move to the affected area. There they remove bacteria and cell debris by phagocytosis. They also release signaling molecules (also produced by macrophages and certain other cells in the immune system) that mediate inflammation.

Review ■

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What are the functions of (1) dendrites, (2) synaptic terminals, and (3) the myelin sheath? How is a neuron different from a nerve? What are the functions of astrocytes? of microglia?

41.3 TRANSMITTING INFORMATION ALONG THE NEURON ■ ■ LEARNING OBJECTIVES 4 Explain how the neuron develops and maintains a resting potential. 5 Compare a graded potential with an action potential, describing the production and transmission of each.

6 Contrast continuous conduction with saltatory conduction.

Most animal cells have a difference in electric charge across the plasma membrane—the electric charge inside the cell is more negative than the electric charge of the extracellular fluid. The plasma membrane is said to be electrically polarized, meaning that one side, or pole, has a different charge than the other side. When electric charges are separated in this way, a potential energy difference exists across the membrane. The difference in electric charge across the plasma membrane gives rise to an electrical gradient. Voltage is the force that causes charged particles to flow between two points. The voltage measured across the plasma membrane is called the membrane potential. If the charges are permitted to come together, they have the ability to do work. Thus, the cell can be thought of as a biological battery. In excitable cells, such as neurons and muscle cells, the membrane potential can change rapidly. Such changes can transmit signals to other cells.

Ion channels and pumps maintain the resting potential of the neuron The membrane potential in a resting (not excited) neuron or muscle cell is its resting potential. The resting potential is generally expressed in units called millivolts (mV). (A millivolt equals one thousandth of a volt.) Like other cells that can produce electrical signals, the neuron has a resting potential of about 70 mV. By convention this is expressed as −70 mV because the cytosol close to the plasma membrane is negatively charged relative to the extracellular fluid (FIG. 41-4a). Biologists can measure the potential across the membrane by placing one electrode inside the cell and a second electrode outside the cell and connecting them through a very sensitive voltmeter or oscilloscope. If you place both electrodes on the outside surface of the neuron, no potential difference between them is registered. (All points on the same side of the membrane have the same charge.) However, once one of the electrodes penetrates the cell, the voltage changes from zero to approximately −70 mV. Two main factors determine the magnitude of the membrane potential: (1) differences in the concentrations between specific ions inside the cell and those in the extracellular fluid and (2) selective permeability of the plasma membrane to these ions. The distribution of ions inside neurons and in the extracellular fluid surrounding them is like that of most other cells in the body. The potassium ion (K+) concentration is about 10 times greater inside the cell than outside the cell. In contrast, the sodium ion (Na+) concentration is about 10 times greater outside than inside. This asymmetric distribution of ions across the plasma membrane at

rest is brought about by the action of selective ion channels and ion pumps. In vertebrate neurons (and skeletal muscle fibers), the resting membrane potential depends mainly on the diffusion of ions down their concentration gradients (see discussion of diffusion in Chapter 5).

Axon

40 20 0 –20 –40 –60

–70 mV

–80

Time

Amplifier Plasma membrane

Electrode placed inside the cell

+

+

–

+

–

– +

– +

Electrode placed outside the cell

+

–

–

– +

– +

+

+

–

–

– +

– +

(a) Measuring the resting potential of a neuron. The axon is negatively charged compared with the surrounding extracellular fluid. The difference in electric charge across the plasma membrane is measured by placing an electrode just inside the neuron and a second electrode in the extracellular fluid just outside the plasma membrane. Extracellular fluid

+

K+

+

–

– +

K+

K

K+

Cl–

+

+

–

–

K+

+

+

–

–

Na+ Cl–

Na+

Na+

Na+

Diffusion in

Na+/K+ pump

Cl–

A–

Cytoplasm

Cl–

Plasma membrane

K+

Na+

A– Cl–

Na

K+

Diffusion out K+

+

3 Na+ Na+

Cl–

K+ 2 K+

A–

(b) Permeability of the neuron membrane. The membrane is less permeable to Na+ than to K+. Passive ion channels allow K+ to diffuse out along their concentration gradient. Potassium ions are mainly responsible for the voltage across the membrane. Sodium–potassium pumps in the plasma membrane actively pump Na+ out of the cell and pump K+ in. The plasma membrane is permeable to negatively charged Cl–, and they contribute slightly to the negative charge. Large anions (A–) such as proteins also contribute to the negative charge.

FIGURE 41-4 Animated Resting potential

Ions cross the plasma membrane by diffusion through ion channels Recall that net movement of ions occurs from an area of higher concentration of that type of ion to one of lower concentration. Membrane proteins form ion channels that typically allow only specific types of ions to pass. Neurons have three types of ion channels: passive ion channels, voltage-activated channels, and chemically activated ion channels. Passive ion channels permit the passage of specific ions such as Na+, K+, Cl−, and Ca2+ (FIG. 41-4b). Unlike voltage-activated and chemically activated ion channels, passive ion channels are not controlled by gates. Potassium channels are the most common type of passive ion channel in the plasma membrane, and cells are more permeable to potassium than to other ions. In fact, in the resting neuron, the plasma membrane is up to 100 times more permeable to K+ than to Na+. When pumped out of the neuron, Na+ cannot easily pass back into the cell. However, K+ pumped into the neuron easily diffuses out. Potassium ions leak out through passive ion channels following their concentration gradient. As K+ diffuse out of the neuron, they increase the positive charge in the extracellular fluid relative to the charge inside the cell. The resulting change in the electrical gradient across the membrane influences the flow of ions. This electrical gradient forces some of the positively charged potassium ions back into the cell. The membrane potential at which the flux (flow) of K+ inward (due to the electrical gradient) equals the flux of K+ outward (because of the concentration gradient) is the equilibrium potential for K+. The equilibrium potential for any particular ion is a steady state in which opposing chemical and electrical fluxes are equal and there is no net movement of the ion. The equilibrium potential for K+ in the typical neuron is −80 mV. The equilibrium potential for Na+ is −40 mV; it differs from the equilibrium potential of K+ because of the difference in concentration of Na+ across the membrane (concentration is high outside and low inside the cell). Because the membrane is much more permeable to K+ than to Na+, the resting potential of the neuron is closer to the potassium equilibrium potential than to the sodium equilibrium potential. (Remember that the resting potential of the neuron is about −70 mV.) The resting potential is primarily established by the net flow of K+ into the neuron. However, the net flow of Na+ outward contributes to the resting potential. Because the plasma membrane is permeable to negatively charged chloride ions, Cl− also contribute slightly. These ions accumulate in the cytosol close to the plasma membrane. Negative charges on large molecules such as proteins add to the negative charge in the cytosol. These large anions cannot cross the plasma membrane. Ion pumping maintains the gradients that determine the resting potential The neuron plasma membrane has very efficient sodium– potassium pumps that actively transport Na+ out of the cell and

K+ into the cell (see Fig. 41-4b; also see Fig. 5-17). These pumps require ATP to pump Na+ and K+ against their concentration and electrical gradients. For every three Na+ pumped out of the cell, two K+ are pumped in. Thus, more positive ions are pumped out than in. The sodium–potassium pumps maintain a higher concentration of K+ inside the cell than outside, and a higher concentration of Na+ outside than inside. In summary, the development of the resting potential depends mainly on the diffusion of K+ out of the cell. However, once the neuron reaches a steady state, net Na+ diffusion into the cell is greater than net K+ diffusion out of the cell. This situation is offset by the sodium–potassium pumps that pump three Na+ out of the cell for every two K+ pumped in.

Graded local signals vary in magnitude Neurons are excitable cells. They respond to stimuli and can convert stimuli into nerve impulses. An electrical, chemical, or mechanical stimulus may alter the resting potential by increasing the membrane’s permeability to sodium ions. When a stimulus causes the membrane potential to become less negative (closer to zero) than the resting potential, that region of the membrane is depolarized. Because depolarization brings a neuron closer to transmitting a neural impulse, it is described as excitatory. In contrast, when the membrane potential becomes more negative than the resting potential, the membrane is hyperpolarized. Hyperpolarization is inhibitory; it decreases the ability of the neuron to generate a neural impulse. A stimulus may change the potential in a relatively small region of the plasma membrane. Such a graded potential is a local response. It functions as a signal only over a very short distance because it fades out within a few millimeters of its point of origin. A graded potential varies in magnitude; that is, the potential charge varies depending on the strength of the stimulus applied. The larger the stimulus is, the larger the change in permeability and the greater the change in membrane potential.

Axons transmit signals called action potentials When a stimulus is strong enough, a rapid, large change in membrane potential occurs, depolarizing the membrane to a critical point known as the threshold level. At that point, the neuron fires a nerve impulse, or action potential, an electrical signal that travels rapidly down the axon into the synaptic terminals. All cells can generate graded potentials, but only neurons, muscle cells, and a few cells of the endocrine and immune systems can generate action potentials. In a series of experiments in the 1940s, pioneering English researchers Alan Hodgkin and Andrew Huxley inserted electrodes into large axons found in squids. By measuring voltage changes as they varied ion concentrations, these investigators showed that passage of Na+ ions into the neuron and K+ ions out of the neuron resulted in an action potential. The process of Na+ rushing into the cell can be compared to the gas pedal of the action, whereas the process of K+ leaving the cell brakes the action potential. In 1963, Hodgkin and Huxley received the Nobel Prize in Physiology or Medicine for this research.

We now know that changes in voltage regulate specific voltageactivated ion channels (also called voltage-gated ion channels) in the plasma membrane of the axon and cell body. Biologists can study these channels by using the patch clamp technique to measure currents across a very small segment of the plasma membrane rather than across the entire membrane (FIG. 41-5). The patch of membrane measured is so small that it may contain a single channel. Biologists have used certain toxins that affect the nervous system to investigate how membrane channels function. For example, the poison tetrodotoxin (TTX) binds to voltage-activated Na+ channels, blocking the passage of Na+. Using TTX and other toxins, researchers were able to identify the channel protein. TTX was first isolated from the Japanese puffer fish, which is eaten as a delicacy (FIG. 41-6). In small amounts TTX tingles the taste buds, but in a larger dose it can prevent breathing. (The Japanese government operates a certification program for puffer fish chefs!) Investigators have established that voltage-activated ion channels have charged regions that act as gates. Voltage-activated Na+ channels have two gates—an activation gate and an inactivation gate (FIG. 41-7a). Voltage-activated K+ channels have a single gate (FIG. 41-7b). Local anesthetics such as procaine (novocaine) and lidocaine (Xylocaine), as well as cocaine, bind to voltage-activated Na+ channels and block them. The channels cannot open in response to depolarization, and the neuron cannot transmit an impulse through the anesthetized region. Signals do not reach the brain, so pain is not experienced. An action potential is generated when the voltage reaches threshold level The membrane of most neurons can depolarize by about 15 mV, that is, to a potential of about −55 mV, without initiating an action potential. However, when depolarization is greater than −55 mV, the threshold level is reached and an action potential is generated. The neuron membrane quickly reaches zero potential and even overshoots to +35 mV or more as a momentary reversal in polarity takes place. The sharp rise and fall of the action potential is called a spike. FIGURE 41-8a on page 870 illustrates an action potential recorded by placing one electrode inside an axon and one just outside. At resting potentials, activation gates of the voltage-activated Na+ channels are closed and Na+ cannot pass into the neuron (FIG. 41-8b; follow the steps illustrated in this figure as you read about transmission of action potentials, repolarization, and return to the resting state). When the voltage reaches threshold level, the channel proteins change shape, opening the activation gates. Sodium ions can then flow through the voltage-activated Na+ channels. As a result, the inside of the neuron becomes positively charged relative to the fluid outside the plasma membrane, generating an action potential. Inactivation gates that are open in the resting neuron close slowly in response to depolarization. Sodium ions can pass through the channel only during the brief period when both activation and inactivation gates are open. The generation of an action potential depends on a positive feedback system. (Recall from Chapter 39 that in a positive feedback system, a change in some condition triggers a response that intensifies the change.) As just discussed, when a neuron is

RESEARCH METHOD Why Is It Used?

The patch clamp technique allows researchers to study the action of a single ion channel over time, even in very small cells. This research method was developed in the mid-1970s by Erwin Neher and Bert Sakmann, both of the Max Planck Institute in Germany. They were awarded a Nobel Prize in 1991 for this work. Plasma membrane

Micropipette Ion channel Ion channel

Cell

Micropipette

How Is It Done?

The tip of a micropipette is tightly sealed to a patch of membrane so small that it generally contains only a single ion channel. The flow of ions through the channel can be measured using an extremely sensitive recording device. Cell biologists have found that the current flow is intermittent and corresponds to the opening and closing of the ion channel. The permeability of the channel affects the magnitude of the current.

FIGURE 41-5 The patch clamp technique The patch clamp technique has been modified in many ways and has been applied to studies of the roles of ion channels in a wide range of cellular processes in both plants and animals. For example, until recently neuroscientists thought that transmitting action potentials was a very inefficient process. They thought that there was a high degree of overlap between the time that Na+ rush into the cell and K+ move out of the cell. This would be a waste of energy because it would be like stepping on the gas and the brake at the same time. Henrik Alle, a neu-

depolarized, voltage-activated Na+ channels open, increasing the membrane permeability to Na+. Sodium ions diffuse into the cell, moving from an area of higher concentration to an area of lower concentration. As Na+ flow into the cell, the cytosol becomes positively charged relative to the extracellular fluid. This charge further depolarizes the neuron so that more voltage-activated Na+ channels open, which increases membrane permeability to Na+. The neuron repolarizes and returns to a resting state After a certain period, inactivation gates close in the Na+ channels, and the membrane again becomes impermeable to Na+. This

roscientist at the Max Planck Institute for Brain Research in Frankfurt, Germany, and his colleagues recently used the patch clamp technique to record currents in neurons of the learning and memory areas of rat brains. They found that the flow of Na+ and of K+ were almost completely separated in time. Na+ have almost finished entering the neuron before K+ ions begin to exit. In contrast to earlier estimates of 30% efficiency for transmission of signals along the neuron, neurons may transmit action potentials with more than 70% efficiency.

inactivation begins the process of repolarization, during which the membrane potential returns to its resting level. Voltageactivated K+ channels slowly open in response to depolarization. When Na+ have almost stopped rushing into the neuron, K+ channels open and K+ leak out of the neuron. This decrease in intracellular K+ returns the interior of the membrane to its relatively negative state, repolarizing the membrane. Voltage-activated K+ channels remain open until the resting potential has been restored. As a wave of depolarization moves down the membrane of the neuron, the normal polarized state is quickly re-established behind

refractory periods, most neurons can transmit several hundred impulses per second.

Jeffrey L. Rotman/Peter Arnold, Inc.

In summary, neural action proceeds as follows:

FIGURE 41-6 Japanese puffer fish A poison, tetrodotoxin, isolated from this fish (Fugu rubripes) binds to voltageactivated sodium ion channels, allowing neurobiologists to study them.

it. The entire process—depolarization and then repolarization— can take place in less than 1 millisecond. During the millisecond or so in which it is depolarized, the axon membrane is in an absolute refractory period: it cannot transmit another action potential no matter how great a stimulus is applied because the voltage-activated Na+ channels are inactivated. Until their gates are reset, they cannot open again. When enough Na+ channel gates have been reset, the neuron enters a relative refractory period that lasts for a few more milliseconds. During this period, the axon can transmit impulses, but the threshold is higher (less negative). Even with the limits imposed by their

Extracellular fluid

neuron in resting state ¡ stimulus causes depolarization ¡ threshold reached ¡ action potential transmits signal ¡ repolarization and return to resting state

The action potential is an all-or-none response Any stimulus too weak to depolarize the plasma membrane to threshold level cannot fire the neuron. It merely sets up a local signal that fades and dies within a few millimeters from the point of stimulus. Only a stimulus strong enough to depolarize the membrane to its critical threshold level results in transmission of an impulse along the axon. An action potential is an all-or-none response because either it occurs or it does not. No variation exists in the strength of a single impulse. If the all-or-none law is valid, how can we explain differences in the levels of intensity of sensations? After all, we have no difficulty distinguishing between the pain of a severe toothache and that of a minor cut on the arm. We can explain this apparent inconsistency by understanding that the intensity of sensation depends on several factors, including the number of neurons stimulated and on their frequency of discharge. For example, suppose you burn your hand. If a large area is burned, more pain receptors are stimulated and more neurons become depolarized. Also, each neuron transmits more action potentials per unit of time.

+ Na+ Na

Na+

Na+

K+

Na+

K+

–70mV

–55mV

Activation gate

Na+ Na+

Cytoplasm

K+

K+

Na+

Inactivation gate

(a) Sodium channels. In the resting state, voltageactivated Na+ channels are closed (left). When the voltage reaches threshold level, activation gates open quickly, allowing Na+ to pass into the cell (right). After a certain time, inactivation gates close, blocking the channels (not shown). Inactivation gates are open when a neuron is in the resting state; they close slowly in response to depolarization. The ion channel is open only during the brief period when both activation and inactivation gates are open.

FIGURE 41-7 Voltage-activated ion channels

K+

K+

K+

K+

(b) Potassium channels. Voltageactivated K+ channels have activation gates that open slowly in response to depolarization. The activation gates close after the resting potential has been restored. Potassium channels have no inactivation gates.

When the neuron plasma membrane is depolarized to threshold level, voltage-activated Na+ channels open and Na+ rush in, causing greater depolarization; an action potential is generated.

KEY POINT

40

Spike

Membrane potential (mV)

20

Depolarization

Repolarization

–20 Threshold level

–40 –60

Axon

3

2

0

–80

Resting state

4

1

0

1

2 3 4 5 Time (milliseconds)

6

7

(a) Action potential. When the axon depolarizes to about –55 mV, an action potential is generated. (The numerical values vary for different nerve cells.) Extracellular fluid +

Na

Na+

Na+ Na+

K+

Na+

Na+

Na+

K+

K+

K+

Na+

Na+

Na+

Na+

+

+

+

+

–

–

–

–

–

–

–

–

–

+

+

+

+

Sodium channel

Na+ +

K

Cytoplasm 1

K+

K+ K+

Potassium channel

Na+

+

K+

Resting state. Voltageactivated Na+ and K+ channels are closed.

Na

Na+

K+

2

K+

Na+

K+

K+

K

K+

+

+

+

+

+

+

+

–

–

–

–

–

–

–

Na+

Na+

K+

Depolarization. At threshold, voltage-activated Na+ channels open. Na+ rush into the neuron, causing further depolarization. Action potential is generated.

3

K+ Na+ Na+

K+

Na+ +

Na+

Na Na+

K+

K+

K+

+

Na+

Na

K+

Na+

K+

K+

+

Na+ +

K+

–

Na+

Na+ Na+

Na+

K+ K+

K+

+

Na+

K+

Na+

K+

+

+

–

–

– Na+

Na+

+

4 Repolarization. Voltageactivated Na+ channels become inactivated. Voltageactivated K+ channels are open; K+ diffuse out of cell, restoring negative charge to inside of cell.

Na+

Na+

K+

+

Na+

Na

Na+

K+

Na+

+

K+

K+

K+

Na

K+

Na+

Na+

Return to resting state. Voltage-activated Na+ and K+ channels close. Na+/K+ pumps (not shown) move Na+ out of the cell and K+ into the cell, restoring the original ion distribution.

(b) The action of ion channels in the plasma membrane determines the state of the neuron.

FIGURE 41-8 Animated Neural signaling along a neuron Generation of an action potential depends on the action of ion channels in the plasma membrane.

An action potential is self-propagating Once initiated, an action potential continues along the neuron. During depolarization, the affected area of the membrane is more positive relative to adjacent regions where the membrane is still at resting potential. The difference in potentials between active and resting membrane regions causes ions to flow between them (an electric current). The flow of Na+ into the adjacent region induces the opening of voltage-activated Na+ channels in that area of the

membrane (FIG. 41-9). The process is repeated like a chain reaction until the end of the axon is reached. Thus, an action potential is a wave of depolarization that travels down the length of the axon. Transmission of an action potential is sometimes compared with a flame moving along a trail of gunpowder. Once the gunpowder is ignited at one end of the trail, the flame moves steadily to the other end by igniting the powder particles ahead of it. An amazing difference is that, after a brief refractory period, the neuron recovers and is able to transmit another action potential.

Stimulus

Axon

Area of depolarization

Na+

K+

+ –

+

–

–

+

+

+

+

+

+

+

+

+

–

–

–

–

–

–

– – Sodium channel

–

Na+

Na+

+

+

Na+

–

–

+

+

+

+

–

–

–

–

–

–

–

–

–

+

+

–

–

–

–

+

+

+

+

+

+

+

+

+

Na+

Na+

Na+

Action potential is transmitted as wave of depolarization that travels down axon. At region of depolarization, Na+ diffuse into cell. Area of repolarization

K+

+ –

+ –

K+

+ –

+

– + K+

2

Potassium channel + +

–

Action potential Na+

K+

1

Na+

–

Na+

K+ Na+ K+

Na+

–

+ –

K+

+

+

–

–

K

+

K

K

K+

K+

K+

– +

– + K+

– +

– + K+

Area of depolarization

+

+

+

–

–

–

+

–

Na+

K+

K

+

+

Na

+

–

+

+

+

–

Na+

+

– +

–

+

+

+

–

–

Na+

Action potential Na+

–

K+

Na+

–

Na+

+

K+

Na+

–

Na+

+ –

Na+

+

+

+

–

–

–

–

–

+

+

Na+

Na+

As action potential progresses along axon, repolarization occurs quickly behind it.

FIGURE 41-9 Transmission of an action potential along the axon When the dendrites (or cell body) of a neuron are stimulated sufficiently to depolarize the membrane to its threshold level, a strong pulse of voltage change travels down the axon. This pulse is an action potential.

Several factors determine the velocity of an action potential Compared with the flow of electrons in electrical wiring or the speed of light, a nerve impulse travels rather slowly. Most axons transmit impulses at about 1 to 10 m per second. The smooth, progressive impulse transmission just described, called continuous conduction, occurs in unmyelinated neurons. In these axons, the speed of transmission is proportional to the diameter of the axon.

Axons with larger diameters transmit faster than do those with smaller diameters because an axon with a large diameter presents less internal resistance to the ions flowing along its length. Squids and certain other invertebrates have giant axons, up to 1 mm in diameter, which allow them to respond rapidly when escaping from predators. In vertebrates, another strategy has evolved that speeds transmission—myelinated neurons. Myelin acts as an effective electrical insulator around the axon. However, the axon is not myelinated at the nodes of Ranvier. At these nodes, the plasma membrane of the axon makes direct contact with the surrounding extracellular fluid. Voltage-activated Na+ and K+ channels are concentrated at the nodes. In fact, in myelinated neurons, ion movement across the membrane occurs only at the nodes. The ion activity at an active node results in diffusion of ions along the axon that depolarize the next node. The action potential appears to jump from one node of Ranvier to the next (FIG. 41-10). This type of impulse transmission is known as saltatory conduction (from the Latin word saltus, which means “to leap”). In myelinated neurons, the distance between successive nodes of Ranvier affects speed of transmission. When nodes are farther apart, less of the axon must depolarize and the axon conducts the impulse faster. Using saltatory conduction, a myelinated axon can conduct an impulse up to 50 times faster than the fastest unmyelinated axon. Saltatory conduction has another advantage over continuous conduction: it requires less energy. Ions move across the plasma membrane only at the nodes, so fewer Na+ and K+ are displaced. As a result, the sodium–potassium pumps do not expend as much ATP to re-establish resting conditions each time an impulse is conducted.

Review ■

■

■ ■ ■

How do ion channels and sodium–potassium pumps contribute to the resting potential? What sequence of events occurs when threshold level is reached in a neuron? How do voltage-activated ion channels work? How does an action potential differ from a graded potential? What are the benefits of saltatory conduction compared with continuous conduction?

KEY POINT

In saltatory conduction, Na+ rapidly diffuse from one node of Ranvier to the next in a myelinated neuron. Action potentials appear to leap from node to node.

neuron that terminates at a specific synapse is called a presynaptic neuron, whereas a neuron that begins at that synapse is a postsynaptic neuron. Note that these terms are relative to a specific synapse. A neuron that is postsynaptic with respect to one synapse may be presynaptic to the next synapse in the sequence.

Signals across synapses can be electrical or chemical

Area of action potential

Saltatory conduction

Nodes of Ranvier

– + + –

+ – – +

+ – – +

+ – – +

– + + –

+ – – +

+ – – +

+ – – +

+ – – +

– + + –

+ – – +

+ – – +

+ – – +

+ – – +

– + + –

1

+ – – + Axon Schwann cell

2

3

4

Direction of depolarization

FIGURE 41-10 Animated Saltatory conduction This sequence of diagrams (steps 1 through 4) illustrates transmission of an action potential in a myelinated neuron. The arrows in the axons indicate diffusion of Na+.

41.4 TRANSMITTING INFORMATION ACROSS SYNAPSES ■ ■ LEARNING OBJECTIVES 7 8 9

Describe the actions of the neurotransmitters identified in the chapter. Trace the events that take place in synaptic transmission and draw diagrams to support your description. Compare excitatory and inhibitory signals and their effects.

A synapse is a junction between two neurons or between a neuron and an effector, such as between a neuron and a muscle cell. A

Based on how presynaptic and postsynaptic neurons communicate, two types of synapses have been identified: electrical synapses and chemical synapses. In electrical synapses, the presynaptic and postsynaptic neurons occur very close together (within 2 nm of one another) and form gap junctions (see Chapter 5). The interiors of the two cells are physically connected by a protein channel. Electrical synapses let ions pass from one cell to another, permitting an impulse to be directly and rapidly transmitted from a presynaptic to a postsynaptic neuron. The escape responses of many animals involve electrical synapses. For example, the “tail-flick” escape response of the crayfish involves giant neurons in the nerve cord that form electrical synapses with large motor neurons. The motor neurons signal muscles in the abdomen to contract. Electrical synapses are important in the mammalian nervous system as well. For example, many interneurons communicate by way of electrical synapses. The majority of synapses are chemical synapses. Presynaptic and postsynaptic neurons are separated by a space, the synaptic cleft, about 20 nm wide (less than one-millionth of an inch). Because depolarization is a property of the plasma membrane, when an action potential reaches the end of the axon, it cannot jump the gap. The electrical signal must be converted into a chemical one. Neurotransmitters are the chemical messengers that conduct the neural signal across the synapse and bind to chemically activated ion channels in the membrane of the postsynaptic neuron. This binding triggers specific gated ion channels to open (or close), resulting in changes in permeability of the postsynaptic membrane. When a postsynaptic neuron reaches its threshold level of depolarization, it transmits an action potential.

Neurons use neurotransmitters to signal other cells Many chemical compounds are now known or suspected to function as neurotransmitters, chemical messengers used by neurons to signal other neurons or to signal muscle or gland cells. We can categorize neurotransmitters into several chemical groups, including acetylcholine, biogenic amines, amino acids, neuropeptides, and gaseous neurotransmitters (TABLE 41-2). Acetylcholine is a low-molecular-weight neurotransmitter that is released from motor neurons and triggers muscle contraction. Acetylcholine is also released by some neurons in the brain and in the autonomic nervous system (see Inquiring About: Alzheimer’s Disease; also see Chapter 42). Cells that release acetylcholine are called cholinergic neurons. Neurons that release norepinephrine are called adrenergic neurons. Norepinephrine, epinephrine, and the neurotrans-

TABLE 41-2

Selected Neurotransmitters

Neurotransmitter

Structure

Site of Action

Action

CNS, neuromuscular junctions

Excitatory effect on skeletal muscle; inhibitory effect on cardiac muscle; excitatory or inhibitory at other sites; acetylcholine concentration in the brain decreases during the progression of Alzheimer’s disease

CNS, PNS

Excitatory or inhibitory; concentration in brain affects mood, sleep, wakefulness, attention

CNS

Helps maintain balance between excitation and inhibition of neurons; important in motor functions; concentration in brain affects mood; also involved in motivation and reward

CNS

Excitatory effect on pathways that control muscle action; inhibitory effect on sensory pathways; helps regulate food intake, sleep, and wakefulness; affects mood

CH2 COOH

CNS, invertebrate neuromuscular junctions

Principal excitatory neurotransmitter in vertebrate brain; functions in learning and memory; excitatory effect at invertebrate neuromuscular junctions

COOH

CNS

Excitatory; functions in learning and memory

O

Acetylcholine

H2C C

Biogenic Amines Norepinephrine

CH2 CH2 N+

O

[CH3]3

HO CH

HO

CH2

NH2

OH HO

Dopamine

CH2 CH2

HO Serotonin (5-hydroxytryptamine, 5-HT)

HO

C CH2 N H

Amino Acids Glutamate

H2N

CH

NH2

CH2 NH2

CH

CH2

COOH

Aspartate

H2N

CH

CH2

COOH Glycine

H2N

CH2 COOH

CNS

Inhibitory (mainly in spinal cord)

Gamma-aminobutyric acid (GABA)

H2N

CH2 CH2

CNS, invertebrate neuromuscular junctions

Major inhibitory neurotransmitter in mammalian brain

CNS, PNS

Endogenous opiates; bind with opiate receptors on afferent fibers that transmit pain signals; inhibit release of substance P

R¬P¬K¬P¬Q¬Q¬F¬F¬G¬L¬M

CNS, PNS

Activates ascending pathways that transmit pain signals

NO

Most neurons?

Retrograde messenger; transmits signals from postsynaptic to presynaptic neuron

CO

Most neurons?

May be neuromodulator

Neuropeptides Endorphins and enkephalins

Substance P Gaseous Neurotransmitters Nitric oxide

Carbon monoxide

CH2 COOH

Y¬G¬G¬F¬M* (Methionine enkephalin)

*Letters represent amino acids; refer to Figure 3-17.

mitter dopamine are catecholamines. The catecholamines and the neurotransmitters serotonin and histamine belong to a class of compounds called biogenic amines. Biogenic amines affect

mood, and their imbalance has been linked to several disorders, including major depression, anxiety disorders, attention deficit hyperactivity disorder (ADHD), and schizophrenia.

In qu i ri n g A b o u t ALZHEIMER’S DISEASE peptide appears to form the plaques. Investigators have identified mutations in specific genes that result in the production of abnormal amyloid-b. Just how the longer amyloid-b leads to AD is not known. The abnormal peptide appears to disrupt calcium regulation, killing neurons in the brain. The peptide may also damage mitochondria, releasing harmful free radicals. AD appears to begin with subtle changes in synaptic function in the hippocampus, a region of the brain important in memory. An early symptom is decreased ability to encode new memories. The loss of synaptic function may be caused by abnormal amyloid-b. Mutations in two other proteins, presenillin 1 and presenillin 2, have been linked to AD. These proteins also reduce the ability to regulate calcium concentration. (They affect Ca+ release from the ER.) Recent studies have linked amyloid-b with sleep deprivation. Short-term sleep deprivation (three weeks) apparently increases deposition of amyloid plaque in mouse models. Another group of investigators have demonstrated that amyloid-b can destroy certain types of bacteria, and therefore may have an immune function. Neurofibrillary tangles, which form in the neuron cytoplasm, are another piece of the AD puzzle. The cytoskeletal protein tau normally associates with the protein tubulin, which forms microtubules. Mutations in the tau gene may interfere with the way the tau protein binds to microtubules, resulting in abnormal microtubules that cannot effectively transport materials through axons.

Abnormal tau proteins accumulate in the cytoplasm, forming fibrous deposits that make up the neurofibrillary tangles (see figure). These tangles interfere with neural signaling. Mechanisms that cause amyloid plaques and neurofibrillary tangles are connected. Using transgenic mice, researchers have demonstrated that amyloid-b deposits in the brain are responsible for the formation of neurofibrillary tangles in areas of the brain known to be affected in Alzheimer’s disease. The drug memantine (Namenda), one of the few drugs approved for treating AD, slows cognitive decline but does not prevent the eventual damage that occurs in the brain. Memantine blocks one type of glutamate receptor (NMDA receptor), which appears to help restore calcium balance. Donepezil (Aricept) inhibits acetylcholinesterase so that it does not break down acetylcholine as rapidly. Acetylcholine is a neurotransmitter in the brain important for memory and thought. Several other drugs designed to slow the progress of Alzheimer’s disease are in clinical trials. One of the challenges of developing a drug has been getting the medication through the blood–brain barrier. Some drugs block formation of b-amyloid, whereas others dissolve the abnormal b-amyloid deposits that are already present. Immune mechanisms that target b-amyloid are being investigated. Anti-inflammatory drugs are also being tested as strategies that may prevent or slow the development of AD.

Amyloid plaques

Neurofibrillary tangles

Brain tissue from the cerebral cortex of a patient with Alzheimer’s disease, showing characteristic amyloid plaques (rounded brown areas) and neurofibrillary tangles (dark teardrop shapes).

Thomas Deerinck, NCMIR/Photo Researchers, Inc.

What is Alzheimer’s disease and what causes it? Alzheimer’s disease (AD) is a progressive, degenerative brain disorder in which neural signaling is disrupted and neurons become damaged and die prematurely. Alzheimer’s disease is the leading cause of senile dementia, the loss of memory, judgment, and ability to reason that is sometimes associated with aging. AD can affect the ability to recognize family members and to carry on routine daily activities, such as dressing, bathing, and eating. More than five million people in the United States have AD, and it is one of the top ten leading causes of death. The development of AD may be a lifelong process. One startling study demonstrated that a person’s writing early in life can predict the disease. Young adults whose writings had a lower density of ideas were more likely to develop the disease. One approach to detecting development of AD is the use of positron emission tomography (PET) scans to study glucose uptake in the brain. Low glucose uptake in certain areas of the brain suggests that neurons in these areas may be damaged. More recently, new genetic markers for AD have been identified. In AD, neurons are damaged in certain parts of the brain, including the cerebral cortex and hippocampus, areas that are important in thinking and remembering. Neurons that secrete the neurotransmitter acetylcholine are especially affected. Investigators have demonstrated that two of the abnormalities that develop in brain tissue as we age, amyloid plaques and neurofibrillary tangles, are especially characteristic of AD. Understanding the biochemistry and genetic basis of both plaques and tangles may provide clues to the causes and cures of AD. Amyloid plaques were first identified in autopsied brains in 1906 by the German physician Alois Alzheimer. These plaques are extracellular deposits of a peptide called amyloid-beta (amyloid-b). Neurons near these plaques appear swollen and misshapen. Microglia are typically present, suggesting an attempt to remove the damaged cells or the plaques themselves. Researchers have found that amyloid-b is cut from a precursor protein called b-APP, a large transmembrane protein coded by a gene on chromosome 21. Normal amyloid-b molecules have 40 amino acids. However, about 10% of the peptides produced have two additional amino acids. This longer

Several amino acids function as neurotransmitters. Glutamate is the major excitatory neurotransmitter in the brain. One type of glutamate receptor is the target of several mind-altering drugs, including phencyclidine (“angel dust”). The amino acid glycine and a modified amino acid, gammaaminobutyric acid (GABA), are neurotransmitters that inhibit neurons in the spinal cord and brain. Glycine is the main inhibitory neurotransmitter in the spinal cord. GABA is the neurotransmitter used at most rapid inhibitory synapses throughout the brain. Many drugs that reduce anxiety enhance the actions of GABA by permitting lower concentrations of GABA to open Cl− channels. These drugs include benzodiazepines, such as diazepam (Valium) and alprazolam (Xanax), and barbiturates, such as phenobarbital. Neuropeptides, such as endorphins and enkephalins, appear to function mainly as neuromodulators, messenger molecules that stimulate long-term changes that enhance or inhibit synaptic function. Some neurobiologists consider them a group of neurotransmitters, but others classify them as a separate group. Endorphins, such as beta-endorphin, and enkephalins are the body’s own endogenous opioids. Opiate drugs, for example, morphine and codeine, are powerful analgesics, drugs that relieve pain without causing loss of consciousness. Endorphins and enkephalins bind to opioid receptors (the same receptors to which opiate drugs bind) and block pain signals. Opioids modulate the effect of other neurotransmitters, including substance P, which activates pathways that transmit pain signals from certain sensory neurons to the CNS. (We discuss pain perception in Chapter 43.) The signaling molecule nitric oxide (NO) is a gas that acts as a retrograde messenger at some synapses. It transmits information from the postsynaptic neuron to the presynaptic neuron, the opposite direction of transmission from other neurotransmitters. Carbon monoxide has also been identified as a gaseous neurotransmitter that may function as a neuromodulator. Some neurobiologists consider nitric oxide and carbon monoxide to be second messengers rather than neurotransmitters.

Neurotransmitters bind with receptors on postsynaptic cells Neurotransmitters are stored in synaptic terminals within hundreds of small membrane-enclosed sacs called synaptic vesicles (FIG. 41-11a). Each time an action potential reaches a synaptic terminal, voltage-gated Ca2+ channels open. Calcium ions from the extracellular fluid then flow into the synaptic terminal. The Ca2+ induce synaptic vesicles to fuse with the presynaptic membrane and release neurotransmitter molecules into the synaptic cleft by exocytosis (FIG. 41-11b). Neurotransmitter molecules diffuse across the synaptic cleft and bind with specific receptors on the dendrites or cell bodies of postsynaptic neurons (or on the plasma membranes of effector cells). Many neurotransmitter receptors are chemically activated ion channels known as ligand-gated ion channels. (See Chapter 6.) When the neurotransmitter, the ligand, binds with the receptor, the ion channel opens (FIG. 41-11c). (Recall that a ligand is a molecule that binds to a specific site in a receptor.) The acetyl-

choline receptor, for example, is a ligand-gated ion channel that permits the passage of Na+ and K+. Some neurotransmitters, such as serotonin, operate by a different mechanism. They work indirectly through a second messenger. Binding of the neurotransmitter with a receptor activates a G protein. The G protein then activates an enzyme, such as adenylyl cyclase, in the postsynaptic membrane. Adenylyl cyclase converts ATP to cyclic AMP (cAMP), which acts as a second messenger (see Chapters 6 and 49). Cyclic AMP activates a kinase that phosphorylates a protein, which then closes K+ channels. For a postsynaptic neuron to repolarize quickly, any excess neurotransmitter in the synaptic cleft must be removed. Some neurotransmitters are inactivated by enzymes. For example, excess acetylcholine is degraded into its chemical components, choline and acetate, by the enzyme acetylcholinesterase. Other neurotransmitters, such as the biogenic amines, are actively transported back into the synaptic terminals, a process known as reuptake. These neurotransmitters are repackaged in vesicles and recycled. Many drugs inhibit the reuptake of neurotransmitters. For example, some antidepressants are selective serotonin reuptake inhibitors (SSRIs). SSRIs such as fluoxetine (Prozac) and escitalopram (Lexapro) selectively inhibit the reuptake of serotonin in the brain. This action concentrates serotonin in the synaptic cleft, elevating mood. Some antidepressants (and cocaine) also inhibit the reuptake of dopamine.

Activated receptors can send excitatory or inhibitory signals A postsynaptic neuron may have receptors for several types of neurotransmitters. Some of its receptors may be excitatory, and others may be inhibitory. Depending on the type of postsynaptic receptor with which it combines, the same neurotransmitter can have different effects. For example, acetylcholine has an excitatory effect on skeletal muscle and an inhibitory effect on cardiac muscle. When neurotransmitter molecules bind to receptors, the activated receptor can directly or indirectly open or close ion channels; this action changes the permeability of the postsynaptic membrane to certain ions. The resulting redistribution of ions changes the electrical potential of the membrane, and the membrane may depolarize. For example, when neurotransmitter molecules combine with receptors that open Na+ channels, the resulting influx of Na+ partially depolarizes the membrane. If sufficiently intense, a local depolarization can set off an action potential. A change in membrane potential that brings the neuron closer to firing is called an excitatory postsynaptic potential (EPSP). Imagine that sufficient sodium ions enter to change the membrane potential from −70 mV to −60 mV. The membrane would be only −5 mV away from threshold. Under such conditions, an additional relatively weak stimulus, causing a small influx of Na+, can cause the neuron to fire. Note that unlike action potentials, postsynaptic potentials are graded responses. Some neurotransmitter–receptor combinations hyperpolarize the postsynaptic membrane. The membrane potential becomes more negative. Because such an action takes the neuron farther

KEY POINT

At most synapses the action potential cannot jump across the synaptic cleft between the two neurons. This problem is solved by chemical signaling across synapses.

Synaptic vesicles

J. F. Gennaro/Photo Researchers, Inc.

Axon of presynaptic neuron Voltage-gated Ca2+ channel

Synaptic terminal

1

Ca2+

Plasma membrane of presynaptic neuron Plasma membrane of postsynaptic neuron

0.25 μm

Synaptic vesicles

2

(a) The TEM shows synaptic terminals filled with synaptic vesicles. Neurotransmitter molecule

3

4

Ligand-gated channel Postsynaptic neuron

5

Ca2+

Postsynaptic membrane

Synaptic terminal

Receptor for neurotransmitter

(b) How a neural impulse is transmitted across a synapse. 1

Action potential reaches synaptic terminals at end of presynaptic neuron.

2

Calcium channels open in membrane, letting Ca2+ from extracellular fluid enter the synaptic terminal.

3

Ca2+ cause synaptic vesicles to fuse with plasma membrane and release neurotransmitter into synaptic cleft.

4

Neurotransmitter binds with receptors on membrane of postsynaptic neuron.

5

In response, specific ion channels open or close, resulting in either depolarization or hyperpolarization. When depolarization reaches threshold level, an action potential is generated in postsynaptic neuron.

Synaptic cleft

Presynaptic membrane Na+

Postsynaptic membrane

(c) Neurotransmitter binds with receptor. Ligand-gated channel opens, resulting in depolarization.

FIGURE 41-11 Animated Neural signaling across a synapse

away from the firing level, a potential change in this direction is called an inhibitory postsynaptic potential (IPSP). For example, if the membrane potential changes from −70 mV to −80 mV, the membrane is farther away from threshold and a stronger stimulus is required to fire the neuron. Like an EPSP, an IPSP can be produced in several ways. Two types of GABA receptors are known to produce IPSPs. When GABA binds to a GABAB receptor, K+ channels open. As K+ diffuse out, the neuron becomes more negative, hyperpolarizing the membrane. Activated GABA A receptors produce IPSPs by opening Cl− channels. In this case, the influx of negative ions hyperpolarizes the membrane. Although the mechanisms are different, the inhibiting effect of GABA is the same in both cases. In summary, neural signaling across synapses involves the following sequence of events: action potential reaches synaptic terminals ¡ calcium ions enter synaptic terminal ¡ synaptic vesicles release neurotransmitter into the synaptic cleft ¡ neurotransmitter diffuses across synaptic cleft ¡ neurotransmitter binds with receptors in membrane of postsynaptic neuron ¡ ion channels open or close, resulting in depolarization or hyperpolarization of postsynaptic membrane

Review ■ ■

What are some functions of biogenic amines? of GABA? What events take place at a chemical synapse? How are EPSPs produced? IPSPs?

41.5 NEURAL INTEGRATION ■ ■ LEARNING OBJECTIVE 10 Define neural integration and describe how a postsynaptic neuron integrates incoming stimuli and “decides” whether to fire.

Each neuron may synapse with hundreds of other neurons. Indeed, thousands of synaptic terminals of presynaptic neurons may cover as much as 40% of a postsynaptic neuron’s dendritic surface and cell body. It is the job of the dendrites and the cell body of every neuron to integrate the messages that continually bombard them. Neural integration is the process of summing, or integrating, incoming signals. EPSPs and IPSPs are produced continually in postsynaptic

Postsynaptic membrane potential (mV)

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neurons, and IPSPs cancel the effects of some EPSPs. It is important to remember that each EPSP and IPSP is not an all-or-none response. Rather, each is a graded response that does not travel like an action potential but may be added to or subtracted from other EPSPs and IPSPs. As the postsynaptic neuron membrane continuously updates its molecular tabulations, the neuron may be inhibited or brought to threshold level. This mechanism integrates hundreds of signals (EPSPs and IPSPs) before an action potential is actually transmitted along the axon of a postsynaptic neuron. Local responses permit the neuron and the entire nervous system a far greater range of response than would be possible if every EPSP generated an action potential.

Postsynaptic potentials are summed over time and space Each EPSP or IPSP is a graded potential that varies in magnitude depending on the strength of the stimulus applied. One EPSP is usually too weak to trigger an action potential by itself. Its effect is below threshold level. However, even though subthreshold EPSPs do not produce an action potential, they do affect the membrane potential. EPSPs can be added together in a process called summation (FIG. 41-12). Temporal summation occurs when repeated stimuli cause new EPSPs to develop before previous EPSPs have decayed. The summation of several EPSPs may bring the neuron to the critical firing level.

+30 Threshold level

0

Resting potential −50 −70 E1

E1

E1 E1

E1

E2 E1 + E2

E1

I

E1 + I

Time (msec)

(a) Subthreshold (no summation). Each stimulus is at a subthreshold level, so no action potential is generated.

(b) Temporal summation. The second EPSP occurs before the first dies off, so the two EPSPs are summed. An action potential is generated.

(c) Spatial summation. EPSPs generated by input from two different presynaptic neurons are summed.

FIGURE 41-12 Neural integration E1 and E2 are excitatory stimuli. I represents an inhibitory stimulus.

(d) Spatial summation of EPSPs and IPSPs. EPSPs and IPSPs are also summed and can cancel each other out.

Spatial summation can also bring the postsynaptic neuron to threshold level. Spatial summation occurs when several closely spaced synaptic terminals release neurotransmitter simultaneously. The postsynaptic neuron is stimulated at several places at once. Neurons also sum IPSPs. As IPSPs are added together as a result of temporal and spatial summation, they move the membrane potential farther away from threshold.

Where does neural integration take place? On a molecular level, every neuron sorts through the thousands of bits of information simultaneously bombarding it. In vertebrates more than 90% of the neurons in the body are located in the CNS, so most neural integration takes place there, within the brain and spinal cord. These neurons are responsible for making most of the “decisions.” In the next chapter, the brain and spinal cord are examined in some detail.

Review ■

What is neural integration? How do temporal and spatial summation differ?

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Where does most neural integration take place in vertebrates?

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(a) Convergence of neural input. Several presynaptic neurons synapse with one postsynaptic neuron.

41.6 NEURAL CIRCUITS: COMPLEX INFORMATION SIGNALING ■ ■ LEARNING OBJECTIVE 11 Distinguish between convergence and divergence, and explain why each is important.

The CNS contains millions of neurons, but it is not just a tangled mass of nerve cells. Its neurons are organized into neural networks, and within each network the neurons are arranged in specific pathways, or neural circuits. Although each network has some special characteristics, the neural circuits in all the networks share many organizational features. For example, convergence and divergence are probably characteristic of them all. In convergence, a single neuron is controlled by converging signals from two or more presynaptic neurons (FIG. 41-13a). An interneuron in the spinal cord, for instance, may receive converging information from sensory neurons entering the cord, from neurons bringing information from various parts of the brain, and from neurons coming from different levels of the spinal cord. Information from all these sources must be integrated before an action potential is transmitted and an appropriate motor neuron is stimulated. Convergence is an important mechanism by which the CNS integrates the information that impinges on it from various sources. In divergence, a single presynaptic neuron stimulates many postsynaptic neurons (FIG. 41-13b). Each presynaptic neuron may branch and synapse with thousands of different postsynaptic neurons. For example, a single neuron transmitting an impulse from the motor area of the brain may synapse with hundreds of interneurons in the spinal cord, and each of these may diverge, in turn, so that hundreds of muscle fibers are stimulated.

(b) Divergence of neural output. A single presynaptic neuron synapses with many postsynaptic neurons.

Review ■

FIGURE 41-13 Neural circuits

■ ■

How does convergence differ from divergence?

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

41.1 (page 861) 1 Describe the processes involved in neural signaling: reception, transmission, integration, and action by effectors. Neural signaling involves (1) reception of information by a sensory receptor; (2) transmission by an afferent neuron to the central nervous system (CNS), which consists of the brain and spinal cord; (3) integration by interneurons in the central nervous system; (4) transmission by an efferent neuron to other neurons or to an effector; and (5) action by effectors, the muscles and glands.

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Sensory receptors and neurons outside the CNS make up the peripheral nervous system (PNS).

41.2 (page 862) 2 Describe the structure of a typical neuron and give the function of each of its parts. ■ Neurons are specialized to receive stimuli and transmit electrical and chemical signals. In a typical neuron, a cell body contains the nucleus and most of the organelles. Many branched dendrites

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extend from the cell body; they are specialized to receive stimuli and send signals to the cell body. The single long axon extends from the cell body and forms branches called axon collaterals. The axon transmits signals into its terminal branches, which end in synaptic terminals. Many axons are surrounded by an insulating myelin sheath. In the PNS the myelin sheath is formed by Schwann cells. In the CNS the sheath is formed by other glial cells. Nodes of Ranvier are gaps in the sheath between successive Schwann cells. A nerve consists of several hundred axons wrapped in connective tissue; a ganglion is a mass of neuron cell bodies in the PNS.

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potential becomes more negative than the resting potential, the membrane is hyperpolarized. A graded potential is a local response that varies in magnitude depending on the strength of the applied stimulus. A graded potential fades out within a few millimeters of its point of origin. If voltage across the membrane declines to a critical threshold level, voltage-activated ion channels open, allowing Na+ to flow into the neuron, and an action potential is generated. The action potential is a wave of depolarization that moves down the axon. As the action potential moves down the axon, repolarization occurs behind it. During depolarization, the axon enters an absolute refractory period, a time when it cannot transmit another action potential. When enough gates controlling Na+ channels have been reset, the neuron enters a relative refractory period, a time when the threshold to fire the neuron is higher. Learn more about ion channels and action potentials by clicking on the figures in CengageNOW.

The action potential conforms to an all-or-none response; no variation exists in the strength of a single impulse. Either the membrane potential exceeds threshold level, leading to transmission of an action potential, or it does not. Once begun, an action potential is self-propagating. 6 Contrast continuous conduction with saltatory conduction. ■ Continuous conduction, which involves the entire axon plasma membrane, takes place in unmyelinated neurons. ■ Saltatory conduction, which is more rapid than continuous conduction, takes place in myelinated neurons. Depolarization appears to skip along the axon from one node of Ranvier to the next; these are sites where the axon is not covered by myelin and where Na+ channels are concentrated. ■

Learn more about the structure of neurons and nerves by clicking on the figures in CengageNOW.

3 Name the main types of glial cells and describe the functions of each. ■

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Glial cells support and nourish neurons and are important in neural communication. Astrocytes are glial cells that physically support neurons. They help regulate the composition of the extracellular fluid in the CNS by taking up excess potassium ions. Astrocytes communicate with one another and with neurons; they also induce and stabilize synapses. Oligodendrocytes are glial cells that form myelin sheaths around axons in the CNS; Schwann cells form sheaths around axons in the PNS. Ependymal cells line cavities in the CNS and help produce and circulate cerebrospinal fluid. They may also serve as neural stem cells. Microglia are phagocytic cells.

41.3 (page 865) 4 Explain how the neuron develops and maintains a resting potential. Neurons use electrical signals to transmit information along axons. The plasma membrane of a resting neuron, one that is not transmitting an impulse, is polarized. The inner surface of the plasma membrane is negatively charged relative to the extracellular fluid just outside the membrane. ■ The resting potential is the potential difference of about −70 mV that exists across the membrane. The magnitude of the resting potential is determined by (1) differences in concentrations of specific ions (mainly Na+ and K+) inside the cell relative to the extracellular fluid and (2) selective permeability of the plasma membrane to these ions. ■ Ions pass through specific passive ion channels. K+ leak out more readily than Na+ can leak in. The resting potential is mainly due to the net flow of K+ into the neuron. Cl− and large anions inside the neuron contribute negative charges. ■ The gradients that determine the resting potential are maintained by ATP-requiring sodium–potassium pumps that continuously transport three Na+ out of the neuron for every two K+ transported in. 5 Compare a graded potential with an action potential, describing the production and transmission of each. ■ If a stimulus causes the membrane potential to become less negative, the membrane becomes depolarized. If the membrane ■

41.4 (page 872) 7 Describe the actions of the neurotransmitters identified in the chapter. The junction between two neurons or between a neuron and effector is a synapse. Although there are electrical synapses, most synapses are chemical. In synaptic transmission, a presynaptic neuron releases neurotransmitter, a chemical messenger, from its synaptic vesicles. ■ Acetylcholine triggers contraction of skeletal muscle. ■ The biogenic amines include norepinephrine, epinephrine, serotonin, dopamine, and histamine. These neurotransmitters play important roles in regulating mood. Dopamine is also important in motor function. Several amino acids function as neurotransmitters, including glu■ tamate, the main excitatory neurotransmitter in the brain, and GABA, a widespread inhibitory neurotransmitter. ■ The neuropeptides, which are opioids, include the endorphins (for example, beta-endorphin) and the enkephalins. Nitric oxide (NO), a gaseous neurotransmitter, transmits signals ■ from a postsynaptic neuron to a presynaptic neuron, the opposite direction from most neurotransmitters. 8 Trace the events that take place in synaptic transmission and draw diagrams to support your description. Calcium ions induce synaptic vesicles to fuse with the presynaptic ■ membrane and release neurotransmitter into the synaptic cleft. The neurotransmitter diffuses across the synaptic cleft and combines with specific receptors on a postsynaptic neuron. Many neurotransmitter receptors are proteins that form ligand■ gated ion channels. Others activate a G protein and work through a second messenger such as cAMP. ■

Learn more about synaptic transmission by clicking on the figure in CengageNOW.

9 Compare excitatory and inhibitory signals and their effects. ■

Binding of a neurotransmitter to a receptor can cause either an excitatory postsynaptic potential (EPSP) or an inhibitory postsynaptic potential (IPSP), depending on the type of receptor. EPSPs bring the neuron closer to firing. IPSPs move the neuron farther away from its firing level.

41.5 (page 877) 10 Define neural integration and describe how a postsynaptic neuron integrates incoming stimuli and “decides” whether to fire. ■ Neural integration is the process of summing, or integrating, incoming signals. ■ Each EPSP or IPSP is a graded potential that varies in magnitude depending on the strength of the stimulus applied. Summation is the process of adding and subtracting incoming signals. By summation of several EPSPs, the neuron may be brought to critical firing level.

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Temporal summation occurs when repeated stimuli cause new EPSPs to develop before previous EPSPs have decayed. Spatial summation occurs when several closely spaced synaptic terminals release neurotransmitter simultaneously, stimulating the postsynaptic neuron at several different places.

41.6 (page 878) 11 Distinguish between convergence and divergence, and explain why each is important. ■ Complex neural circuits are possible because of associations such as convergence and divergence. In convergence, a single neuron is affected by converging signals from two or more presynaptic neurons. Convergence allows the CNS to integrate incoming information from various sources. In divergence, a single presynaptic neuron stimulates many postsynaptic neurons, allowing widespread effect.

T E S T YO U R U N D E R S TA N D I N G 1. Summing incoming neural signals is part of (a) reception (b) transmission (c) integration (d) action by effectors (e) afferent neuron transmission 2. Action potentials are transmitted to synaptic terminals by the (a) ganglia (b) axon (c) dendrites (d) cell body (e) nodes 3. The myelin sheath is produced around axons in the PNS by (a) axons (b) neuron cell bodies (c) dendrites (d) Schwann cells (e) oligodendrocytes 4. Which of the following does not contribute to the resting potential of a neuron? (a) sodium–potassium pumps (b) ion channels (c) differences in concentration of ions across the membrane (d) differences in permeability to certain ions (e) graded potentials

10. A presynaptic neuron in the cerebrum transmits information to hundreds of other neurons. This is an example of (a) convergence (b) divergence (c) temporal summation (d) spatial summation (e) a graded potential 11. In spatial summation (a) repeated stimuli cause new EPSPs to develop before previous ones decay (b) divergence occurs at two or more synapses (c) EPSPs and IPSPs cancel each other in the space of the axon (d) two or more EPSPs transmit a summed chemical signal (e) several closely spaced synaptic terminals release neurotransmitter simultaneously 12. Describe the action taking place at each of the numbered steps. Use Figure 41-11c to check your answers.

5. Which of the following occurs first when voltage reaches the threshold level? (a) gates of certain voltage-activated ion channels open (b) K+ channels close (c) the membrane hyperpolarizes (d) neurotransmitter is released (e) neurotransmitter reuptake takes place at the synapse 6. Saltatory conduction (a) requires more energy than continuous conduction (b) occurs in unmyelinated neurons (c) occurs when the action potential jumps from one node of Ranvier to the next (d) slows transmission of an impulse (e) depends on the action of GABA 7. Acetylcholine (a) is a biogenic amine (b) is recycled (c) is released by motor neurons and by some neurons in the brain (d) is deactivated by G proteins (e) binds to opioid receptors 8. Receptors for serotonin and some other neurotransmitters (a) are voltage-activated ion channels (b) permit influx of chloride ions, leading to depolarization of the membrane (c) work through a second messenger (d) inhibit reuptake of the neurotransmitter (e) are passive ion channels 9. IPSPs (a) excite presynaptic neurons (b) excite postsynaptic neurons (c) can cancel the effects of some EPSPs (d) release large amounts of neurotransmitters (e) are released by postsynaptic cell bodies

Ca2+

Synaptic terminal

1 2

Synaptic cleft

Presynaptic membrane Na+ 3

Postsynaptic membrane 4

CRITICAL THINKING 1. Discuss several specific ways in which the nervous system helps maintain homeostasis. 2. Stimulant drugs such as amphetamines increase the activity of the nervous system. Describe two mechanisms involving synaptic transmission that could explain the action of such drugs. 3. Develop a hypothesis to explain the fact that acetylcholine has an excitatory effect on skeletal muscle but an inhibitory effect on cardiac muscle. 4. EVOLUTION LINK. From the perspective of adaptation, hypothesize the advantages of neural circuits characterized by a great deal of convergence and divergence. 5. ANALYZING DATA. State whether each of the following would result in an EPSP or IPSP (explain your answer): (a) Na+ enter postsynaptic neuron; (b) K+ diffuse out of postsynaptic neuron; (c) Cl− enter postsynaptic neuron.

6. SCIENCE, TECHNOLOGY, AND SOCIETY. A test for AD risk is now on the market. Based on gene markers that can be identified in a saliva sample, a healthy person can learn his or her approximate risk for AD. Do you think that this information will be useful to people? Would you worry about the psychological consequences of a person knowing that he or she had a high risk? Would you want to know? 7. SCIENCE, TECHNOLOGY, AND SOCIETY. Investigators have genetically engineered cells to produce acetylcholine, dopamine, GABA, and other neurotransmitters. In what ways might these cells be useful in neurobiological research? How might they be useful in clinical medicine? Additional questions are available in CengageNOW at www.cengage.com/ login.

42

Neural Regulation

Volker Steger/Peter Arnold, Inc.

The brain during speech. In this functional magnetic resonance image (fMRI) of the brain, red indicates areas of greatest activation during speech; yellow indicates medium activation.

KEY CONCEPTS 42.1 Trends in invertebrate nervous system evolution include increased number and concentration of neurons and interneurons, more complex synaptic contacts, specialization of function, and cephalization. 42.2 The vertebrate nervous system consists of a central

A

frog flicks out its tongue with lightning speed to capture a fly, a rabbit escapes from a predator, and you learn biology—perhaps not that

quickly. In a complex nervous system, millions of neurons work together to transmit information from the external environment to the central nervous system (CNS). After the information is integrated, neural signals are transmitted to the effectors that can produce effective responses. The nervous system also regulates heart rate, breathing, and hun-

nervous system (CNS), which includes the brain and spinal cord and a peripheral nervous system (PNS), which includes sensory receptors and nerves.

dreds of other internal activities. Nerves must transmit information from

42.3 Although different regions of the brain are special-

the information that will allow organs to make the adjustments neces-

ized in various vertebrate classes, the brain develops in the same way and has the same basic structure; during the evolution of the vertebrate brain, the cerebrum and cerebellum have become larger and more complex.

sary to maintain homeostasis. Researchers have focused extensively on

42.4 As in other vertebrates, the human CNS receives sensory information from both the internal and external environment, integrates it, and determines appropriate responses. The association areas of the brain are responsible for learning, memory, language, thought, and judgment.

42.5 The peripheral nervous system has two divisions: the somatic division transmits information between the external environment and the CNS, whereas the autonomic division transmits information between the internal environment and the CNS.

42.6 Many prescribed and illicit drugs affect mood by affecting neurotransmitter concentrations in the brain.

all of the organs in the body to the CNS. Then, other nerves transmit back

just how neurons interact, and we are only beginning to understand the mechanisms that permit the many vital functions of the nervous system. Neurobiologists use a variety of methods to study the mechanisms of neural function. For example, improved imaging methods have revolutionized the study of the brain. Functional magnetic resonance imaging (fMRI) has provided investigators with a window through which to observe brain function. Functional MRI allows neurobiologists to study the responses of neural networks in the brain while an individual is actually performing a task, such as speaking (see photograph). During the performance of the task, an area of the brain becomes active and flow of oxygenated blood to that area increases. Functional MRI detects changes that take place in response to a very brief stimulus. For example, a visual

stimulus lasting only 30 milliseconds stimulates brain activation that can be detected by fMRI. In this chapter we compare several types of nervous systems. We then examine the structure and function of the vertebrate nervous system, with emphasis on the function of the human brain. We explore some of the frontiers of neurobiology, such as mechanisms involved in information processing and drug addiction.

42.1 INVERTEBRATE NERVOUS SYSTEMS: TRENDS IN EVOLUTION ■ ■ LEARNING OBJECTIVES 1 2

Contrast nerve nets and radial nervous systems with bilateral nervous systems. Identify trends in the evolution of the invertebrate nervous system.

An animal’s lifestyle is closely related to the organization and complexity of its nervous system. Hydra and other cnidarians have a nerve net consisting of interconnected neurons with no central control organ. Sensory cells are located in the epidermis, and the cnidarian can respond to stimuli from any direction (see Chapter 31). When a neuron is stimulated, electrical signals are sent from neuron to neuron in all directions (FIG. 42-1). Responses may involve large parts of the body. An advantage of the nerve net is that the cnidarian responds effectively to predator or prey approaching from any direction. Hydra responds by discharging nematocysts (stinging structures) and

coordinating the movements of its tentacles to capture food. Some cnidarians have two or more nerve nets. In some jellyfish, a slowtransmission nerve net coordinates movement of the tentacles, and a second nerve net, which is faster, coordinates swimming. The radial nervous system of the sea star and other echinoderms (see Chapter 32) is a modified nerve net. This system shows some degree of selective organization of neurons into more than a diffuse network. It consists of a nerve ring around the mouth, from which a large radial nerve extends into each arm. Branches of these nerves, which form a network somewhat similar to the nerve net of Hydra, coordinate the animal’s movement. Bilaterally symmetrical animals have a bilateral nervous system. In planarian flatworms, the head region contains concentrations of nerve cells called cerebral ganglia (FIG. 42-2). These serve as a primitive brain and have some control over the rest of the nervous system. Typically, two solid, ventral, longitudinal nerve cords extend from the ganglia to the posterior end of the body. Transverse nerves connect the two nerve cords and connect the brain with the eyespots. This arrangement is called a “ladder-type” nervous system. Annelids and arthropods typically have a solid ventral nerve cord (FIG. 42-3). The cell bodies of many of the neurons are massed into ganglia. Afferent and efferent neurons lie in lateral nerves that link the ganglia with muscles and other body structures. If an earthworm’s brain is removed, the animal can move almost as well as before. However, when it bumps into an obstacle, it persists in a futile effort to move forward rather than turn aside. The earthworm needs its brain in order to respond adaptively to environmental change.

Cerebral ganglia Eyespot

Nerve cords Transverse nerves Nerve net

FIGURE 42-1 Animated Hydra’s nerve net

FIGURE 42-2 Animated Ladder-type nervous system of flatworms

Hydra and other cnidarians have a network of neurons with no central control organ.

Cerebral ganglia in the head region serve as a simple brain. The longitudinal nerve cords are connected by transverse nerves.

Pharynx

Brain

Dorsal blood vessel

Hearts

Ganglia

Ventral nerve cord

Crop

octopus is capable of considerable learning and can be taught difficult tasks. In fact, cephalopods are considered the most intelligent invertebrates. We can identify the following trends in the evolution of nervous systems:

Intestine

Ventral blood vessel

1. Increased number of nerve cells. 2. Concentration of nerve cells. The nerve cells form masses of tissue that become ganglia (concentrations of cell bodies) and brain, and form thick cords of tissue that become nerve cords and nerves. 3. Specialization of function. For example, transmission of nerve impulses in one direction requires both afferent nerves, which conduct impulses toward a central nervous system (CNS), and efferent nerves, which transmit impulses away from the CNS to the effectors (muscles and glands). Certain parts of the CNS become specialized to perform specific functions, and distinct structural and functional regions can be identified. 4. Increased number of interneurons and more complex synaptic contacts. These contacts permit greater integration of incoming messages, provide a greater range of responses, and allow more precise responses. 5. Cephalization, or formation of a head. A bilaterally symmetrical animal generally moves in a forward direction. Concentration of sense organs at the front end of the body lets the animal detect an enemy quickly enough to escape, or to see or smell food in time to capture it. Response can be rapid if short pathways link sense organs to decision-making nerve cells nearby. Therefore, nerve cells are typically concentrated in the head region and organized to form ganglia or a brain.

Lateral nerves

(a) Earthworm nervous system. Annelids have a dorsal anterior brain and one or more ventral nerve cords. Cell bodies of the neurons are located in ganglia that are connected by the ventral nerve cord. Intestine Brain

Tubular heart

Ventral nerve cord

Ganglia

Review

(b) Insect nervous system. The arthropod brain is continuous with the ventral nerve cord. The brain is more specialized than in annelids.

In what ways is the insect nervous system different from the nervous system of the flatworm? ■ What are some trends in the evolution of nervous systems? ■ What are some advantages of cephalization? ■

FIGURE 42-3 Animated The annelid and arthropod nervous systems

The cerebral ganglia of some arthropods differ from those of annelids in that they have specific functional regions. These areas are specialized for integrating information transmitted to the ganglia from sense organs. Mollusks with inactive lifestyles have relatively simple nervous systems with little cephalization and very simple sense organs. Typically, there are two nerve cords; one extends down each side of the body. Several pairs of ganglia are located along the cords. The cephalopod mollusks (squids and octopods) have complex nervous systems that include well-developed sense organs. The cephalopod nervous system is adapted to the active, predatory lifestyle of these animals. Neurons concentrated in ganglia are massed in a ring surrounding the esophagus; they make up a brain that contains about 168 million nerve cells. Like the vertebrate brain, the cephalopod brain has lobes and intricate folds, and particular areas are specialized as centers for specific functions. The

42.2 THE VERTEBRATE NERVOUS SYSTEM: STRUCTURE AND FUNCTION ■ ■ LEARNING OBJECTIVE 3

Describe the two main divisions of the vertebrate nervous system and summarize their functions.

An animal’s range of possible responses depends in large part on how many neurons it has and how they are organized in the nervous system. Some invertebrate nervous systems have only a few hundred neurons. As animal groups evolved, nervous systems became increasingly complex. The human brain has an estimated 100 billion neurons. How neurons are organized is critical in determining learning ability and flexibility of behavior.

Central Nervous System (CNS)

to

C

fr o d

ed

t te

n

n

tr

tr

it t

m ns

tr a

S

CN

CN

to

tion

S

Peripheral Nervous System (PNS)

ma

I n fo r

m

ed

fr o

mi

io

io

it t

ns

at

ed

tr a

at

it t

ion

S

m

sm

mat

rm

an

N

fo

or

sm

Infor

In

S

m

N

I nf

an

C

Brain and spinal cord

Somatic division

Autonomic division

Receptors

Receptors

Afferent (sensory) nerves transmit signals from receptors to CNS.

Afferent (sensory) nerves transmit signals from receptors in internal organs to CNS.

Efferent (motor) nerves transmit signals from CNS to effectors: skeletal muscles and glands.

Efferent nerves

Sympathetic nerves

Parasympathetic nerves

Transmit signals from CNS to effectors: smooth muscle, cardiac muscle, glands

FIGURE 42-4 Animated Organization of the vertebrate nervous system

The vertebrate nervous system has two main divisions: the central nervous system (CNS) and the peripheral nervous system (PNS) (FIG. 42-4). The CNS consists of a highly developed brain that is continuous with the dorsal, tubular spinal cord. Serving as central control, these organs integrate incoming information and determine appropriate responses. The PNS is made up of the sensory receptors (for example, tactile, auditory, and visual receptors) and the nerves, which are the communication lines. Various parts of the body are linked to the brain by cranial nerves and to the spinal cord by spinal nerves. Afferent (sensory) neurons in these nerves continuously inform the CNS of changing conditions. Then efferent neurons transmit the “decisions” of the CNS to appropriate muscles and glands, which make the adjustments needed to maintain homeostasis.

For convenience the PNS is subdivided into somatic and autonomic divisions. Most of the receptors and nerves concerned with changes in the external environment are somatic. Those that regulate the internal environment are autonomic. Both divisions have afferent nerves, which transmit messages from receptors to the CNS, and efferent nerves, which transmit information back from the CNS to the structures that respond. The autonomic division has two kinds of efferent pathways: sympathetic and parasympathetic nerves (discussed later in the chapter).

Review ■ ■

What are the main components of the CNS? What are the main components and functions of the PNS?

42.3 EVOLUTION OF THE VERTEBRATE BRAIN ■ ■ LEARNING OBJECTIVE

Telencephalon Diencephalon

Forebrain

4 Trace the development of the principal vertebrate brain regions from

Mesencephalon

Midbrain

the hindbrain, midbrain, and forebrain, and compare the brains of fishes, amphibians, reptiles, birds, and mammals.

Metencephalon Hindbrain

All vertebrates, from fishes to mammals, have the same basic brain structure, although different parts of the brain are specialized in the various vertebrate classes. The evolutionary trend is toward increasing complexity, especially of the cerebrum and cerebellum. In the early vertebrate embryo, the brain and spinal cord differentiate from a single tube of tissue, the neural tube. Anteriorly, the tube expands and develops into the brain. Posteriorly, the tube becomes the spinal cord. Brain and spinal cord remain continuous, and their cavities communicate. As the brain begins to differentiate, three bulges become visible: the hindbrain, midbrain, and forebrain (FIG. 42-5).

The hindbrain develops into the medulla, pons, and cerebellum As indicated in TABLE 42-1, the hindbrain subdivides to form the metencephalon, which gives rise to the cerebellum and pons, and the myelencephalon, which gives rise to the medulla. The medulla, pons, and midbrain make up the brain stem, the elongated portion of the brain that looks like a stalk holding up the cerebrum. The medulla, the most posterior part of the brain, is continuous with the spinal cord. Its cavity, the fourth ventricle, is continuous with the central canal of the spinal cord and with a channel that runs through the midbrain. The walls of the medulla are thick and made up largely of nerve tracts (bundles of axons) that connect the spinal cord with various parts of the brain. The medulla contains centers that regulate life-sustaining functions such as respiration, heartbeat, and blood pressure. Other reflex centers in the medulla regulate activities such as swallowing, coughing, and vomiting. The cerebellum is responsible for muscle tone, posture, and equilibrium. It also refines and coordinates muscle activity. In hu-

TABLE 42-1 Early Embryonic Divisions

Myelencephalon

FIGURE 42-5 Animated Early development of the vertebrate nervous system Early in the development of the vertebrate embryo, the anterior end of the neural tube differentiates into the hindbrain, midbrain, and forebrain. These primary divisions subdivide and eventually give rise to specific structures of the adult brain.

mans, the cerebellum provides input to motor areas in the cerebral cortex, and in this way helps plan and initiate voluntary activity. Certain regions of the cerebellum store implicit memories (unconscious memories for perceptual and motor skills, such as swimming or skating). The size and shape of the cerebellum vary among the vertebrate classes (FIG. 42-6). Development of the cerebellum is roughly correlated with the extent and complexity of muscular activity, reflecting the principle that the relative size of a brain part correlates with its importance to the behavior of the species. In some fishes, birds, and mammals, the cerebellum is highly developed, whereas it tends to be small in jawless fishes, amphibians, and reptiles. In birds and mammals, the cerebellum is large with many folds. Injury or removal of the cerebellum results in impaired muscle coordination. A bird without a cerebellum cannot fly, and its wings thrash about jerkily. When the human cerebellum is injured by a blow or by disease, muscular movements are uncoordinated. Any activity requiring delicate coordination, such as threading a needle, becomes very difficult, if not impossible.

Differentiation of CNS Structures Subdivisions

Derivatives in Adult

Cavity

Telencephalon

Cerebrum

Lateral ventricles (first and second ventricles)

Diencephalon

Thalamus, hypothalamus, epiphysis (pineal body)

Third ventricle

Midbrain

Mesencephalon

Optic lobes in fish and amphibians; superior and inferior colliculi in mammals

Cerebral aqueduct

Hindbrain

Metencephalon

Cerebellum, pons

Myelencephalon

Medulla

Fourth ventricle

Spinal cord

Central canal

Brain Forebrain

Spinal cord

KEY POINT

Although different parts of the brain are specialized in various vertebrate groups, there is an evolutionary trend toward larger size and greater complexity of the cerebrum and cerebellum. Olfactory bulb Olfactory tract Olfactory lobe Epiphysis Corpus striatum Optic lobe Cerebellum Medulla

(a) Shark

(b) Codfish Olfactory bulb Olfactory tract Epiphysis Cerebrum Diencephalon Optic lobe Cerebellum Medulla

(c) Amphibian (frog)

(d) Reptile (alligator)

Olfactory bulb

Convolutions

Sulcus Cerebrum

FIGURE 42-6 Animated Evolution of the

vertebrate brain

Epiphysis Optic lobe Cerebellum Medulla

(e) Bird (goose)

( f ) Mammal (horse)

Comparison of the brains of members of six vertebrate classes reveals basic similarities and evolutionary trends. (Brains are not drawn to scale.) Note that different parts of the brain are specialized in the various groups. The large olfactory lobes in the shark brain (a) are essential to this predator’s highly developed sense of smell. (b–f) During the course of evolution, the cerebrum and cerebellum have become larger and more complex. In mammals (f), the cerebrum is the most prominent part of the brain; the cerebral cortex, the thin outer layer of the cerebrum, is highly convoluted (folded), which greatly increases its surface area. In (f), the diencephalon and part of the brain stem are covered by the large cerebrum.

In mammals, the pons contains a thick bundle of fibers that transmits information between the two sides of the cerebellum. The pons also serves as a bridge that connects the medulla and cerebellum with other regions of the brain. The pons contains centers that help regulate respiration and nuclei that relay impulses from the cerebrum to the cerebellum. Recall from Chapter 41 that a nucleus is a group of cell bodies within the CNS.

The midbrain is prominent in fishes and amphibians In fishes and amphibians, the midbrain, or mesencephalon, is the most prominent part of the brain and serves as the main association area. It receives incoming sensory information, integrates it, and sends decisions to appropriate motor nerves. The dorsal portion of the midbrain is differentiated to some extent. For example, the optic lobes are specialized for visual interpretations. In reptiles, birds, and mammals, many functions of the optic lobes are assumed by the cerebrum, which develops from the forebrain. In mammals, the midbrain consists of the superior colliculi, centers for visual reflexes such as pupil constriction, and the inferior colliculi, centers for certain auditory reflexes. The inferior colliculi are major integration centers for incoming auditory information. The mammalian midbrain also contains a center (the red nucleus) that helps maintain muscle tone and posture.

The forebrain gives rise to the thalamus, hypothalamus, and cerebrum The forebrain subdivides to form the telencephalon and diencephalon. The diencephalon gives rise to the thalamus and hypothalamus. In all vertebrate classes, the thalamus is a relay center for motor and sensory messages. In mammals, all sensory messages except those from the olfactory receptors are delivered to the thalamus, where they are integrated before they are relayed to the sensory areas of the cerebrum. The hypothalamus, which lies below the thalamus, forms the floor of the third ventricle. The hypothalamus is a major coordinating center for regulating autonomic and somatic responses. It integrates incoming information and provides input to centers in the medulla and spinal cord that regulate activities such as heart rate, respiration, and digestive system function. In birds and mammals, the hypothalamus controls body temperature. It also contains olfactory centers, regulates appetite and water balance, and is important in emotional and sexual responses. As discussed in Chapter 49, the hypothalamus links the nervous and endocrine systems and produces certain hormones. The telencephalon gives rise to the cerebrum. The lateral ventricles (also called the first and second ventricles) lie within the cerebrum. Each lateral ventricle connects with the third ventricle (within the diencephalon) by way of a channel. In most vertebrate groups, the telencephalon also gives rise to the olfactory bulbs. These structures are important in the chemical sense of smell— the dominant sense in most aquatic and terrestrial vertebrates. In fact, much of brain development in vertebrates appears to focus on integrating olfactory information. In fishes and amphibians, a

large part of the cerebrum is devoted to this function. Birds are an exception among the vertebrates in that their sense of smell is generally poor. A part of their cerebrum, the corpus striatum, however, is highly developed. This structure controls eating, flying, singing, and other action patterns. In most vertebrates, the cerebrum is divided into right and left cerebral hemispheres. Most of the cerebrum is made of white matter, which consists mainly of myelinated axons that connect various parts of the brain. In mammals and in the majority of reptiles, a layer of gray matter, the cerebral cortex, makes up the outer portion of the cerebrum. Gray matter is unmyelinated and contains cell bodies and dendrites. Certain reptiles and all mammals have a type of cerebral cortex called the neocortex. Most of the neocortex consists of association areas—regions that link sensory and motor functions and are responsible for higher functions, such as learning and reasoning. The neocortex is very extensive in mammals, making up the bulk of the cerebrum. In humans, most of the cerebral cortex is neocortex, which consists of six distinct cell layers. The remaining part of the cerebral cortex is referred to as the paleocortex. In mammals the paleocortex forms part of the limbic system (discussed later in this chapter). In mammals, the cerebrum is the most prominent part of the brain. During embryonic development, it expands and grows backward, covering many other brain structures. The surface area of the human cerebral cortex is greatly expanded by numerous folds called convolutions. The furrows between them are called sulci (sing., sulcus) if shallow and fissures if deep. The number of convolutions (not the size of the brain) has been associated with complexity of brain function. In mammals, the cerebrum is responsible for numerous functions performed by other parts of the brain in other vertebrates. In addition, the mammalian cerebrum has highly developed association functions.

Review ■

■

What are the structures derived from the vertebrate embryonic forebrain? the hindbrain? What are the functions of these structures? How does the brain of a mammal differ from the brain of an amphibian?

42.4 THE HUMAN CENTRAL NERVOUS SYSTEM ■ ■ LEARNING OBJECTIVES 5 Describe the structure and functions of the human spinal cord. 6 Describe the structure and functions of the human cerebrum. 7 Summarize the sleep–wake cycle and contrast rapid-eye-movement 8

(REM) sleep with non-REM sleep. Describe the actions of the limbic system, including the role of the amygdala in emotional expression.

9 Summarize how the brain processes information, including a description of synaptic plasticity and the neurobiological changes that take place during learning.

The soft, fragile human brain and spinal cord are well-protected. Encased within bone, they are covered by three layers of con-

nective tissue, the meninges. The three meningeal layers are the tough, outer dura mater; the middle arachnoid; and the thin, vascular pia mater, which adheres closely to the tissue of the brain and spinal cord (FIG. 42-7). Meningitis is a disease in which these coverings become infected and inflamed.

Dural sinus (superior sagittal sinus)

The space between the arachnoid and the pia mater is the subarachnoid space, which contains cerebrospinal fluid (CSF). This shock-absorbing fluid cushions the brain and spinal cord against mechanical injury. It also serves as a medium for exchange of nutrients and waste products between the blood and brain. CSF is produced by special networks of capillaries, collectively called the choroid plexus, which extend from the pia mater into the ventricles. After circulating through the ventricles, CSF passes into the subarachnoid space. It is then reabsorbed into large blood sinuses within the dura mater.

Skin of scalp Skull Dura mater Arachnoid Subarachnoid space Pia mater Cortex of brain

The spinal cord transmits impulses to and from the brain

The tubular spinal cord extends from the base of the brain to the level of the second lumbar vertebra. A cross-section view through the spinal cord reveals a small central canal surrounded by an area of gray matter shaped a bit like the letter H (FIG. 42-8). The gray matter consists of large masses of cell bodies, dendrites, unmyelinated axons, and glial cells. (a) Frontal section through the superior part of the brain. Note the dural sinus, a blood sinus, between two layers of The white matter, found outside the the dura mater. Blood leaving the brain flows into such sinuses gray matter, consists of myelinated axand then circulates to the large jugular veins in the neck. ons arranged in bundles called tracts or pathways. Ascending tracts conduct impulses up the cord to the brain. For example, the spinothalamic tracts in the anterior and lateral Skin columns of the white matter conduct pain and temCerebral aqueduct Skull perature information from sensory neurons in the Fourth ventricle skin. The pyramidal tracts are descending tracts Subarachnoid space Pia mater that convey impulses from the cerebrum to motor nerves at various levels in the cord. We describe the Choroid plexus Dura mater spinal nerves later in this chapter. Third ventricle Choroid plexus In addition to transmitting impulses to and from the brain, the spinal cord controls many reflex activities. A reflex action is a relatively simple, involuntary motor response to a stimulus. Although most reflex actions are more complex, let us consider a withdrawal reflex, in which a neural circuit consisting of three types of neurons carries out a response to a stimulus (FIG. 42-9). Suppose that your hand accidentally touches a flame. Almost instantly, and even before you become consciously aware of what has happened, you jerk your hand away. (b) Sagittal section through brain. The CSF, which cushions the brain and The instant your skin contacts the flame, a senspinal cord, is produced by the choroid plexi in the walls of the ventricles. This fluid circulates through the ventricles and subarachnoid space. It is continuously sory neuron transmits a message from pain recepproduced and then reabsorbed into the blood of the dural sinuses. tors in the skin to the spinal cord. Within the spinal cord, a sensory neuron transmits the signal to an interneuron. The interneuron integrates the information and signals FIGURE 42-7 Protection of the brain and spinal cord an appropriate efferent (motor) neuron, which conducts the inforThe CNS is well-protected by the skull and meninges, and by cerebrospinal fluid (CSF). mation to groups of muscles. The muscles respond by contracting,

Dorsal fissure M. I. Walker/Photo Researchers, Inc.

Central canal Gray matter White matter

2.5 mm Ventral root of spinal nerve

(a) Cross section through the spinal cord.

(b) LM of a cross section through the spinal cord.

FIGURE 42-8 Animated Structure of the spinal cord The spinal cord consists of gray matter and white matter.

KEY POINT

A reflex action is a coordinated, involuntary motor response to a stimulus.

Synapse between sensory neuron and interneuron Nerve cell body of sensory neuron

3

1

Receptor

4

5 Muscle

2

Sensory neuron

1

Interneuron

Reception. Sensory receptor signals sensory neuron.

2

FIGURE 42-9 Withdrawal reflex

Transmission. Sensory neuron transmits signal to CNS.

3

Integration. Interneuron integrates information.

4

Nerve cell body of motor neuron

Transmission. Appropriate motor neuron (shown in red ) transmits impulses to muscle.

5

Action by effector. Muscle contracts, moving hand away from flame (actual response).

jerking the hand away from the flame. Actually, many neurons in sensory, association, and motor nerves participate in such a reflex action. Generally, we are not even consciously aware that all these responding muscles exist. In summary, signals are transmitted through a withdrawal reflex in the following sequence: reception by sensory receptor in skin ¡ sensory neuron transmits signal to CNS ¡ interneuron in CNS integrates information ¡ efferent (motor) neuron transmits signal to muscle ¡ muscle contracts

At the same time that the reflex pathway is activated, a message is sent up the spinal cord to the conscious areas of the brain. As you pull your hand away from the flame, you become aware of what has happened and feel the pain. This awareness, however, is not part of the reflex response. The spinal cord has some plasticity and is capable of training. For example, sensory feedback from exercise increases the strength of neural connections in the spinal cord. When descending connec-

TABLE 42-2

tions with the brain are intact, some victims of spinal cord injury (more than 1.2 million people in the United States alone) can develop at least some limited ability to walk again.

The most prominent part of the human brain is the cerebrum The structure and functions of the main parts of the human brain are summarized in TABLE 42-2, and major regions of the brain are illustrated in FIGURES 42-10 and 42-11. As in other mammals, the human cerebral cortex consists of right and left cerebral hemispheres. Specific areas of the cerebrum are specialized for performing specific functions The cerebral cortex is functionally divided into three areas: (1) sensory areas receive incoming signals from the sense organs; (2) motor areas control voluntary movement; and (3) association areas link the sensory and motor areas and are responsible for thought, learning, language, memory, judgment, decision

The Human Brain

Structure

Description

Function

Medulla

Continuous with spinal cord; primarily made up of nerves passing from spinal cord to rest of brain

Contains vital centers (clusters of neuron cell bodies) that control heartbeat, respiration, and blood pressure; contains centers that control swallowing, coughing, vomiting

Pons

Forms bulge on anterior surface of brain stem

Connects various parts of brain with one another; contains respiratory and sleep centers

Midbrain

Just above pons

Center for visual and auditory reflexes (e.g., pupil reflex, blinking, adjusting ear to volume of sound)

Thalamus

At top of brain stem

Main sensory relay center for conducting information between spinal cord and cerebrum; neurons in thalamus sort and interpret all incoming sensory information (except olfaction) before relaying messages to appropriate neurons in cerebrum

Hypothalamus

Just below thalamus; pituitary gland is connected to hypothalamus by stalk of neural tissue

Contains centers for control of body temperature, appetite, and fat metabolism; regulates pituitary gland; important in emotional and sexual responses, and in sleep–wake cycle

Cerebellum

Second-largest division of brain

Muscle coordination and refinement of movements; muscle tone, posture, equilibrium; helps plan and initiate voluntary activity; stores implicit memories

Cerebrum

Largest, most prominent part of human brain; longitudinal fissure divides cerebrum into right and left hemispheres, each divided into lobes: frontal, parietal, temporal, and occipital lobes

Center of intellect, memory, consciousness, and language; also controls sensation and motor functions

Brain stem

Cerebral cortex (outer gray matter)

White matter

Arranged into convolutions (folds) that increase surface area; functionally, cerebral cortex is divided into: 1. Motor areas

1. Control movement of voluntary muscles

2. Sensory areas

2. Receive incoming information from eyes, ears, pressure and touch receptors, etc.

3. Association areas

3. Sites of intellect, memory, language, and emotion; interpret incoming sensory information

Consists of myelinated axons of neurons that connect various regions of brain; axons are arranged into bundles (tracts)

Connects the following: 1. Neurons within same hemisphere 2. Right and left hemispheres 3. Cerebrum with other parts of brain and spinal cord

Parietal lobe

Central sulcus

Frontal lobe

Cerebrum

Prefrontal area

Cerebellum

Fred Hossler/Visuals Unlimited

Occipital lobe

Temporal lobe Brain stem Medulla

(a)

(b)

Brain stem Cerebellum

FIGURE 42-10 Animated The human brain Human brain, lateral view. Each cerebral hemisphere is divided into four major lobes. The diencephalon and part of the brain stem are covered by the cerebrum. Compare the diagram (a) with the photograph of the human brain (b).

making, and personality. You can see positron-emission tomography (PET) scans showing some functional areas of the cerebrum in FIGURE 42-12. Investigators have mapped the cerebral cortex, locating the areas responsible for different functions. The occipital lobes contain the visual centers. Stimulation of these areas, even by a blow on the back of the head, causes the sensation of light; their removal causes blindness. The centers for hearing are located in the temporal lobes of the brain above the ear; stimulation by a blow causes a sensation of noise. Removal of both auditory areas causes deafness. Removal of one does not cause deafness in one ear; instead, it produces a decrease in the auditory acuity of both ears. Association areas in the temporal lobes help us identify incoming stimuli. A groove called the central sulcus crosses the top of each hemisphere from medial to lateral edge. This groove partially separates the frontal lobes from the parietal lobes. The frontal lobes have important motor and association areas. The prefrontal cortex is an association area in each frontal lobe that is crucial in evaluating information, making judgments and decisions, planning, and organizing responses. The motor areas in the frontal lobes control the skeletal muscles. The somatosensory area in the anterior region of the parietal lobes receives information regarding touch, pressure, heat, cold, and pain from sense organs in the skin. This

region also receives information regarding position of the body. Association areas in the parietal lobes are important in helping us pay attention to such incoming information. The size of the motor area in the brain for any given part of the body is proportional to the complexity of movement involved, not to the amount of muscle. Predictably, areas that control the hands and face are relatively large (FIG. 42-13 on page 895). A similar relationship exists between the sensory area and the sensitivity of the region of the skin from which it receives impulses. These relationships are visually represented as the sensory homunculus (“little person”) and the motor homunculus in Figure 42-13. Neural fibers in the brain cross so that one side of the brain controls the opposite side of the body. As a result of another “reversal,” the uppermost part of the cortex controls the lower limbs of the body. When all the areas of known function are plotted, they cover almost all of the rat’s cortex, a large part of the dog’s, a moderate amount of the monkey’s, and only a small part of the total surface of the human cortex. The remaining cortical areas are association areas. Somehow the association regions integrate the diverse impulses reaching the brain so that an appropriate response is made. When disease or accident destroys the functioning of one or more association areas, the ability to recognize certain kinds of symbols

Cerebrum

Corpus callosum Thalamus

Cerebrum

Diencephalon

Corpus callosum Science Pictures Limited/Science Photo Library/Photo Researchers, Inc.

Hypothalamus

Pineal gland

Midbrain

Cerebellum Pons Pituitary gland Medulla

Spinal cord

(a)

(b) Cerebellum

Pons Medulla

FIGURE 42-11 Midsagittal section through the human brain Half of the brain has been cut away, exposing structures normally covered by the cerebrum. Compare the diagram (a) with the photograph of the human brain (b).

may be lost. For example, the names of objects may be forgotten, although their functions are remembered and understood. The two cerebral hemispheres are specialized for certain functions. As will be discussed later in this chapter, the left hemisphere is specialized for language functions. This part of the brain is also specialized for logical decision making and retrieval of facts. The right hemisphere is specialized for emotional processing and for visual-spatial tasks such as recognizing faces. Axons in the white matter of the cerebrum connect parts of the brain The white matter of the cerebrum lies beneath the cerebral cortex. Nerve fibers of the white matter connect the cortical areas with one another and with other parts of the nervous system (see Table 42-2). A large band of white matter, the corpus callosum, connects the right and left hemispheres (see Fig. 42-11). Deep within the white matter of the cerebrum lie the basal ganglia, paired groups of nuclei (gray matter). These nuclei play an important role in coordinating movement. The basal ganglia send signals to and receive signals from the substantia nigra in

the midbrain. Neurons from the substantia nigra that project (extend) to the basal ganglia produce dopamine, a neurotransmitter that helps balance inhibition and excitation of neurons involved in motor function. (See discussion of neurotransmitters in Chapter 41.) Other neurons from the substantia nigra signal nuclei in the thalamus, which in turn relay information to the motor cortex. Neurons from the substantia nigra that signal the thalamus release the neurotransmitter gamma-aminobutyric acid (GABA). Just how these areas work together to coordinate motor function is not yet fully understood. More than one-half million people in the United States alone are affected by Parkinson’s disease, a progressive neurological disorder that affects movement. In this disease, dopamine-producing neurons die over time, decreasing the available dopamine. As a result there is a progressive decrease in effective neural messages regarding motor responses. This results in tremors in the limbs and face, a slowing of movement, poor muscle coordination, and impaired walking and balance. Several medications, including a dopamine precursor (l-DOPA) are used to treat symptoms of the disorder.

Primary motor area (voluntary movement)

Supplementary motor area (on inner surface—not visible; programming of complex movements)

CENTRAL SULCUS

Premotor area (coordination of complex movements)

Somatosensory area (sensations from body surface and awareness of body position) Posterior parietal area (integration of somatosensory and visual input; important for complex movements) Wernicke’s area (speech understanding)

Prefrontal association area (planning for voluntary activity; decision making; personality traits)

PARIETAL LOBE

FRONTAL LOBE

Parietal-temporal-occipital association area (integration of all sensory input; important in language)

Broca’s area (controls ability to speak) Primary auditory area (receives and interprets auditory information)

Limbic association cortex (mostly on inner and bottom surface of temporal lobe; motivation and emotion; memory)

OCCIPITAL LOBE Primary visual area (receives and interprets visual information) TEMPORAL LOBE

CEREBELLUM

BRAIN STEM SPINAL CORD

(a) Various areas of the cerebral cortex are responsible for specific types of neural processing. (Lateral view of the brain).

Courtesy of Washington University School of Medicine, St. Louis

FIGURE 42-12 Functional areas of the cerebrum

The body follows a circadian cycle of sleep and wakefulness Researchers have determined that many animals sleep, including nematodes and fruit flies. Predators typically sleep for longer periods than animals that are preyed upon. For example, rabbits sleep for a few minutes at a time. In some birds and aquatic mammals, one cerebral hemisphere sleeps at a time, allowing the animal to remain aware of its environment. Sleeping with one hemisphere permits eared seals and whales to move to the surface to breathe without awakening. Humans and many other animals follow a 24-hour circadian (daily) cycle of sleeping and then being awake. When we are awake, we are typically aware of the external world and of ourselves, including our thoughts, perceptions, and emotions. In contrast, when we are asleep, the body receives messages from the environment, but we are not aware of them. The suprachiasmatic nucleus, the most important of the body’s biological clocks, is located in the hypothalamus. The suprachiasmatic nucleus receives information about the duration of light and dark from the retina of the eye and transmits information to other nuclei and neurons. The suprachiasmatic nucleus signals

(b) Specific areas of the brain "light up" on positron-emission tomography (PET) scans as an individual performs various tasks. When a specific region of the brain is more active, more blood flows to the area. PET scans detect the magnitude of blood flow. Thus, PET scans are pictures of the brain at work at specific tasks. The color scale at the right indicates progressively more activity from the purple color at the bottom to the white at the top, which indicates maximum activity.

Left hemisphere

Right hemisphere

Front

Frontal lobe

Primary motor area

Top view Central sulcus Somatosensory area

Parietal lobe

Back

Occipital lobe

Top

Top

w ms, and ja Teeth, gu e l gu na Ton mi nx do ary b h a P raInt ans org

ll

M li v Sa Vocalization

Lips

Lower lip

Hip

ee Kn le Ank Toes

Fi ng er s

Le Hip g

Lips

Trunk Shoulder Elbow

Left hemisphere

R M ing In idd Th de le Ne um x Bro ck b Eye w l Face id and eye ba

Wrist

Genitalia

rs

ge

n Fi

ot Fo Toes

d Han tle Lit

Trunk Neck Head Shoulder Arm Elbow rm Foreat is Wr d n Ha le t Lit

R M ing In idd d Th ex le Ey um e b No se Face Upper lip

e gu Ton Sw a at astic llowing ion at ion

Left hemisphere

Jaw

Cross-sectional view

Temporal lobe

(a) Sensory homunculus. This cross section through the anterior region of the parietal lobe shows the distribution of sensory input to the somatosensory area from various parts of the body. The homunculus is proportioned to reflect the relative amount of cerebral cortex that receives sensory information from each body part.

Cross-sectional view

Temporal lobe

(b) Motor homunculus. This cross section through the posterior part of the frontal lobe shows which area of the cerebral cortex controls each body part. Note that more of the cerebral cortex is devoted to controlling those body structures capable of skilled, complex movement. The primary motor area is involved in coordinated movement of many muscle groups.

FIGURE 42-13 Maps of the primary sensory and motor areas

the pineal gland about light and dark. This endocrine gland is located in the midline of the diencephalon. The pineal gland produces melatonin, a hormone that plays a role in regulating the sleep–wake cycle. The pineal gland secretes up to ten times as much melatonin during darkness as during day-

light. Jet lag, the state of fatigue and decreased physical and mental performance we experience when we fly to a different time zone, occurs because the body is no longer synchronized with its light– dark cycle. Some individuals take melatonin supplements to promote sleep, but their effectiveness has not been proven.

The hypothalamus, thalamus, and brain stem are all involved in regulating the sleep–wake cycle. Neurons in these regions secrete several neurotransmitters important in regulating sleep, including acetylcholine, norepinephrine, and serotonin. The reticular activating system (RAS) is a neural pathway within the brain stem and thalamus. The RAS receives messages from neurons in the spinal cord and from many other parts of the nervous system, and communicates with the cerebral cortex by complex neural circuits. When certain neurons of the RAS bombard the cerebral cortex with stimuli, an individual feels alert and can focus on specific thoughts. If the RAS is severely damaged, the person may pass into a deep, permanent coma. After many hours of activity, the sleep–wake cycle may be affected by fatigue of the RA S. Sleep centers are then activated, and their neurons release serotonin. During REM sleep, a stage of sleep characterized by rapid eye movement, neurons in the RAS stimulate heightened activity in certain regions of the brain. After sufficient rest, the inhibitory neurons of the sleep centers become less excitable, and the excitatory neurons of the RAS become more excitable.

can be recorded by a device known as an electroencephalograph. A recording of this electrical activity, called an electroencephalogram (EEG), can be obtained by taping a set of electrodes to standard regions of the scalp and measuring the activity of the cerebral cortex (FIG. 42-14). The EEG shows that the brain is continuously active. Certain patterns of brain waves are elicited by specific types of activity. For example, when you are resting quietly with eyes closed, your brain emits alpha rhythm patterns. In contrast, as you are reading this biology text, your brain is emitting beta rhythm patterns, which have a fast-frequency rhythm. Beta rhythm patterns are characteristic of heightened mental activity, such as information processing. During certain stages of sleep, the brain emits slow, high-amplitude theta waves and delta waves. Certain brain conditions and diseases change the pattern of brain waves. Individuals with epilepsy, for example, exhibit a recognizable, abnormal wave pattern. The location of a brain tumor or the site of brain damage caused by a blow to the head can sometimes be determined by noting the part of the brain that emits abnormal waves.

Specific types of activity are associated with certain brain wave patterns

Sleep progresses through several stages

Brain activity can be studied by measuring and recording the electrical potentials, or “brain waves,” generated by thousands of active neurons in various parts of the brain. This electrical activity

Sleep is an alteration of consciousness during which there is decreased electrical activity in the cerebral cortex and from which a person can be aroused. Based on changes in electrical activity, we can identify four stages of non-REM (nonrapid eye move-

Awake

I Slow-wave

Non-REM sleep, stage IV

Sleep stage

REM

II III IV 0

1

2

3 4 5 Time of sleep (hours)

6

Awake Non-REM sleep

REM sleep

REM sleep

(b) A typical cyclical sleep pattern in a young adult.

Awake, eyes open (a) EEG rhythms during different stages of sleep. During non-REM, stage IV sleep, the brain emits delta waves, slow-frequency, high-amplitude waves. During REM sleep, the brain emits high-frequency activity similar to that of a person who is awake.

FIGURE 42-14 EEG rhythms and cyclical pattern of sleep

7

ment; REM is an acronym for rapid eye movement) sleep during the first hour of sleep. The deepest sleep is thought to occur during the fourth stage, when it is difficult to awaken the sleeping person. Slower-frequency, higher-amplitude waves (theta and delta waves) are characteristic of non-REM sleep. This electrical activity is thought to be generated spontaneously by the cerebral cortex when it is not driven by impulses from other parts of the brain. During non-REM sleep, body movement, heart rate, breathing, blood pressure, metabolic rate, and body temperature all decrease. Every 90 minutes or so, a sleeping person enters the REM sleep stage for a time. During this stage (about 20% of total sleep time), the eyes move about rapidly beneath the closed but fluttering lids. Brain waves change to a desynchronized pattern. Everyone dreams, especially during REM sleep. PET scans of sleeping subjects indicate that during REM sleep, blood flow in the frontal lobes is reduced. In contrast, blood flow increases in limbic system areas that produce visual scenes and emotion. During REM sleep, body movement, heart rate, breathing, blood pressure, metabolic rate, and body temperature all increase. Increased secretion of the neurotransmitter GABA by certain neurons in the brain and spinal cord inhibits muscle contraction in some large groups of muscles. This results in a type of temporary paralysis. Sleep may have several functions Although birds, mammals, and many other animals have a sleep– wake cycle, neurophysiologists are not certain why sleep is necessary. One hypothesis holds that memories are consolidated during REM sleep. Another hypothesis suggests that sleep allows less important synaptic connections set during the day to reset. Non-REM sleep may provide time for the brain to restore itself. When a person stays Frontal lobe awake for unusually long periods, fatigue and irritability result, and even routine tasks are not performed well. For example, fatigue is a leading cause of automobile accidents. Sleep also provides the body with the opportunity to conserve energy and to replace glycogen stores.

Emotions are feeling states that we experience both physiologically and cognitively. Humans are born with the capacity to experience and express emotions, including fear, anger, sadness, and happiness. The amygdala filters incoming sensory information and interprets it in the context of emotional needs and survival. When it perceives danger, the amygdala sends information to other parts of the brain so that appropriate responses can be made. This part of the limbic system is important in the experience of fear and aggression. The amygdala becomes hypersensitive to possible danger following a traumatic experience (see Inquiring About: The Neurobiology of Traumatic Experience). Structures of the limbic system are essential in an infant’s bonding with a caregiver during the first year of life. The development of a secure attachment bond between infant and caregiver is the foundation for all future emotional and social connections between the child and other human beings. The quality of early attachment between child and caregiver affects brain development and future behavior. Healthy attachment allows the baby to express emotions and provides the basis for learning how to regulate emotions. Attachment also determines how vulnerable (or resilient) the individual will be to traumatic experiences later in life. Researchers first discovered in the 1950s that the limbic system is part of an important motivational system when they implanted electrodes in certain areas of the brains of laboratory animals. They found that a rat quickly learns to press a lever that stimulates reward centers in the brain. In fact, the rat will press the lever as

The limbic system affects emotional aspects of behavior The limbic system, present in all mammals, influences the emotional aspects of behavior, evaluates rewards, and is important in motivation. It plays a role in sexual behavior, biological rhythms, and autonomic responses. The limbic system includes regions of the cerebrum (parts of the prefrontal cortex, cingulate cortex, temporal lobe, basal ganglia, the hippocampus, amygdala, and olfactory bulb), regions of the thalamus, hypothalamus, several nuclei in the midbrain, and the neural pathways that connect these structures (FIG. 42-15). The limbic system influences the endocrine system and the autonomic division of the nervous system.

Cingulate cortex Fornix

Thalamus

Hippocampus

Temporal lobe

Amygdala Olfactory bulbs Hypothalamus

FIGURE 42-15 The limbic system The limbic system consists of a ring of forebrain structures that surround the brain stem and are interconnected by complex neural pathways. This system includes the amygdala, hippocampus, parts of the prefrontal cortex, the cingulate cortex, and areas in the thalamus, and hypothalamus. The limbic system is important in evaluating threat, in emotional expression, sexual behavior, motivation, and learning.

In qu i ri n g A b o u t T H E N E U RO B I O LO GY O F T R AU M AT I C E X P E R I E N C E Have you been involved in a serious automobile accident or have you been the victim of a violent crime? Are you a combat veteran of Iraq or Afghanistan? Perhaps you are a survivor of a natural disaster, such as a hurricane or earthquake. Or you may have endured childhood neglect or abuse. Whether you are a survivor of any of these experiences, or of other terrifying events, you may have experienced the effects of trauma. A traumatic experience is an event that causes intense fear, helplessness, or horror and that overwhelms normal coping and defense mechanisms. Most of us have strategies for processing moderately disturbing experiences. For example, if you were involved in a minor car accident, or if you were one of the millions of people who watched by television as people were wounded in Iraq or pulled from the rubble of buildings after earthquakes in Chile or Haiti, you may have been disturbed by the events you experienced or witnessed. You probably processed what you saw and heard by thinking about it and talking to your friends and family about what happened. You may have dreamed about your experience. As the brain actively reviews and sorts out a disturbing experience, we make sense of what happened. The emotional intensity decreases, and we store the memory along with other memories of more ordinary past events. The memory of the uncomfortable experience fades in importance and intensity. Traumatic events are more difficult to process. If you survived an experience in which you thought you were going to die or be seriously hurt or if you witnessed someone else being wounded or killed, your body’s response to real or perceived danger may be very intense. Years later, your brain may still be secreting abnormally high amounts of stress hormones and certain neurotransmitters, and you may remain on red alert, your body ready to deal with any new threat. How does the brain respond in a traumatic experience, and how does the experience affect the brain? When danger threatens, the amygdala sends messages to both the hypothalamus (which signals the autonomic division of the nervous system and the endocrine system) and the neocortex (which allows us to be aware of our experience). The amygdala is programmed to remember the smells, sounds, and sensations that are part of the experience. Until the memories of the experience are fully processed, similar smells, sounds, and

sensations remind us of the traumatic event and trigger the body to prepare for danger. The traumatized individual experiences fear and anxiety. Because they are so overwhelming, traumatic memories are very difficult to process. They cause us so much discomfort and anxiety that we tend to avoid them rather than intentionally focus on them and sort them out. As a result, traumatic memories seem to stay “stuck” in the limbic system. When triggered, the experience may be replayed with its original emotional intensity (a flashback). Some trauma survivors develop Post-traumatic Stress Disorder (PTSD), a condition in which they experience (1) intrusive thoughts, images, sensory experiences, memories, and dreams; (2) an urge to avoid reminders of the traumatic event; and (3) physiological hyperarousal, a condition in which the body remains on high-alert status, continuously scanning the environment for potential danger. Several studies have shown an association between prolonged trauma (for example, severe child abuse) and long-term changes in the brain. These changes include EEG abnormalities, smaller size of certain areas of the brain, compromised right brain development (the right brain is specialized for processing information related to emotion, social interaction, and physiological states); differences in neural circuits connecting cortical and subcortical areas (can decrease the individual’s sense of self and lead to poor connections with other people); abnormal concentrations of certain neurotransmitters (affect mood, ability to inhibit inappropriate behavior); inappropriate modulation of the limbic system (its responses are exaggerated); and long-term changes in the sympathetic nervous system and endocrine response to stress (faster, more intense response to stress). Neuroimaging studies have demonstrated changes in brain function, including over-response by the amygdala and decreased response by the medial prefrontal cortex (which normally inhibits the amygdala). When the amygdala overresponds, the individual experiences anxiety, distress, and physiological hyperarousal. These physiological states can lead to intense emotional responses that are rooted in the traumatic experience. Broca’s area, a region in the posterior part of the left frontal cortex, is also affected by trauma. Broca’s area is critically important in generating words and thus in expressing language (see Fig. 42-12). Re-

searchers have shown that when subjects are exposed to accounts of their traumatic experiences, activity in Broca’s area decreases. This deactivation appears to be the physiological basis for the difficulty that trauma survivors have describing their experience in words. Trauma survivors have described a state of “speechless terror” that they experience during flashbacks of traumatic experiences. Without words, it is difficult to process and resolve a traumatic experience. Based on many studies, it is clear that traumatic experiences can cause both structural and functional changes in the brain. The exaggerated responses of the amygdala to harmless stimuli perceived as threatening are dissociated from cortical (medial prefrontal) activation. Responses generated by the limbic system are rooted in emotion rather than in rationality and judgment (see figure below). PTSD can severely affect and limit one’s life. However, this disorder can be resolved with appropriate treatment.

Reminder of traumatic experience (trigger)

Low response by medial prefrontal cortex (which normally inhibits limbic system) Does not inhibit limbic system

Limbic system (including amygdala) over responds

Distress, anxiety

Hypothalamus

Physiological hyperarousal

Emotional responses rooted in traumatic experience

Some limbic system responses to reminders of traumatic experiences

many as 5000 times per hour, choosing this self-stimulation over food and water, until it drops from exhaustion. We now know that the brain has reward centers that give us pleasure when we carry out vital activities such as eating, drinking, and sexual activity. These centers are important in experiencing emotion and in motivation. When stimulated, they activate reward circuits that allow us to feel pleasure in response to certain experiences. The mesolimbic dopamine pathway, a major reward circuit, includes two important midbrain areas, the substantia nigra and the adjacent ventral tegmental area. These areas, which extend into behavioral control centers in the limbic system, contain the largest group of neurons in the brain that release dopamine. Novel stimuli associated with reward or pleasure activate these dopamine neurons. The neurons signal information about surprising events or stimuli that predict rewards. Such signals may motivate us to act when something important is happening. Dopamine pathways are also important in the neurobiology of drug addiction (discussed later in this chapter). As researchers continue to unravel the mechanisms of dopamine pathways, we may better understand such disorders as attention deficit hyperactivity disorder (ADHD) and schizophrenia, conditions associated with excessive amounts of dopamine and other neurotransmitters in the brain. People with these disorders have difficulty filtering out sensory stimuli. The presence of excess dopamine may drive them to divert their attention to so many sensory stimuli that they have difficulty focusing on the most important stimuli.

Learning and memory involve long-term changes at synapses Neurobiologists are beginning to understand how the nervous system functions, including how we learn and remember. Learning is the process by which we acquire knowledge or skills as a result of experience. For learning to occur, we must be able to remember what we experience. Memory is the process of encoding, storing, and retrieving information or learned skills. An interesting question has been: To what extent are the brain’s functions hard-wired, that is, preset? Many areas of the brain once thought to be hard-wired are now known to be flexible and capable of change. Even animals with very simple nervous systems can learn to repeat behaviors associated with reward and to avoid behaviors that cause pain. Such changes in behavior are possible because of synaptic plasticity, the ability of synaptic connections to change in response to experience. Such change involves structure and function. Familiar examples of plasticity in human motor skills include learning to walk, ride a bicycle, or catch a baseball. At first, you were probably clumsy, but with practice your performance became smoother and more precise. For that to happen, changes must occur at synapses in neural circuits. Similarly, the ability to learn languages, solve problems, and perform scientific experiments depends on synaptic plasticity. What are the mechanisms by which we learn diverse activities, such as skilled movement and the study of biology? Canadian

psychologist Donald O. Hebb proposed in 1949 that when two neurons connected by a synapse (that is, presynaptic and postsynaptic neurons) are active simultaneously, the synapse is strengthened. In 1973, British researchers T.V.P. Bliss and T. J. Lomo found experimental evidence for Hebb’s hypothesis. Investigating rabbit brains, these researchers applied a series of high-frequency electrical stimuli to neurons in the hippocampus, a region that helps convert information into memories. The result was a long-term increase in synaptic strength. We can also credit Hebb for the axiom “Neurons that fire together, wire together.” Experience influences how neural circuits in our brains organize themselves. Learning involves the storage of information and its retrieval Implicit memory, also called procedural memory, is unconscious memory for perceptual and motor skills, such as riding a bicycle. Implicit memory is about “how” to do something and involves repeating the behavior until it becomes routine. Declarative memory, also called explicit memory, involves factual knowledge of people, places, or objects and requires conscious recall of the information. The hippocampus is critical in forming and retrieving declarative memories. How long do you remember? You are constantly bombarded with thousands of bits of sensory information. At this very moment, your eyes are receiving information about the words on this page, the objects around you, and the intensity of the light in the room. At the same time, you may be hearing a variety of sounds— music, your friends talking in the next room, the hum of an air conditioner. Your olfactory epithelium may sense cologne or the smell of coffee. Maybe you are eating as you read. Sensory receptors in your hands may be receiving information regarding the weight and position of your book. We may hold such sensory information in immediate memory for fractions of a second. Most sensory stimuli are not important to remember any longer and are filtered out. Short-term memory lasts only for seconds or for a few minutes. Typically, we can hold only about seven chunks of information (for example, seven words or numbers) at a time in short-term memory. Short-term memory lets us recall information for a few minutes. When we look up a phone number, for example, we usually remember it only long enough to call it. If we need the same number an hour later, most of us have to look it up again. When we select information for long-term memory, we must process it. The brain rehearses the material and encodes it. We recognize patterns and meaningfully associate the stimuli to past experience or knowledge. The hippocampus functions in the formation and retrieval of memories. The hippocampus temporarily holds new information and may integrate various aspects of an experience, including odors, sounds, and other information. It helps place our experiences in categories so that they can be stored along with similar memories. We integrate new information with other knowledge already stored in the brain. Memory consolidation allows memories to be transferred to the cerebral cortex and stored for long periods. Consolidation involves expression of genes and protein synthesis and depends on new neural connections at synapses. Time (several minutes, hours, or even longer) is required for the brain to consolidate a new memory.

If a person suffers a brain concussion, memory of what happened immediately before the incident may be completely lost. This is known as retrograde amnesia. When the hippocampus is damaged, long-term memory may not be impaired, and the patient may be able to recall information stored in the past. However, new short-term memories can no longer be converted to long-term memories. Where are memories stored? When large areas of the mammalian cerebral cortex are destroyed, information is lost somewhat in proportion to the extent of lost tissue. No specific area can be labeled the “memory bank.” Rather, memories seem to be stored within many areas of the brain. For example, visual memories may be stored in the visual centers of the occipital lobes, and auditory memories may be stored in the temporal lobes. Researchers think that memories are integrated in many areas of the brain, including association areas of the cerebral cortex, the amygdala, hippocampus, thalamus, and hypothalamus. Wernicke’s area in the temporal lobe has been identified as an important association area for complex thought processes. Neurons within the association areas form interconnected pathways that permit complicated information transfer. Retrieval of information stored in long-term memory is of considerable interest—especially to students! The challenge is to find information when you need it. When you seem to forget something, the problem may be that you have not searched effectively for the memory. Information retrieval can be improved by careful encoding, for example, by forming strong associations between items. On the other hand, unimportant, unused memories are typically forgotten over time. Forgetting can be useful when it clears the brain of unnecessary information. Neurobiological changes occur during learning Short-term memory involves changes in the neurotransmitter receptors of postsynaptic neurons. These changes strengthen synaptic connections. The receptors are linked by second messengers (for example, cyclic AMP) to ion channels in the plasma membrane. In some types of learning, changes take place in presynaptic terminals or postsynaptic neurons that permanently enhance or inhibit the transmission of impulses. In some cases, specific neurons may become more sensitive to a particular neurotransmitter. Neurons typically transmit action potentials in bursts, and the amount of a neurotransmitter released by each action potential may increase or decrease. Repeated electrical stimulation of neurons causes a functional change at synapses. When a presynaptic neuron continues to transmit action potentials at a high rate for a minute or longer, there is a long-lasting strengthening of the connection between the presynaptic and postsynaptic neurons. This increased strength of the synaptic connection is known as long-term potentiation (LTP). Potentiation means “to strengthen or make more potent.” In contrast, low-frequency stimulation of neurons results in a long-lasting decrease in the strength of their synaptic connections. This decrease is called long-term depression (LTD; not related to the mood disorder). Information storage and forgetting depend on the strengthening and weakening of synaptic connections brought about by LTP and LTD. Induction of LTP and LTD requires activation of two types of glutamate receptors: NMDA receptors and AMPA receptors.

(Each of these receptors is named for the compound that artificially activates it. NMDA is the acronym for N-methyl-D-aspartate and AMPA stands for a-amino-3-hydroxyl-5-methyl-4-isoxazolepropionate.) NMDA and AMPA receptors are present on the plasma membranes of postsynaptic neurons. They control the passage of calcium ions into neurons. NMDA receptors respond to the neurotransmitter glutamate by opening Ca2+ channels. However, when the postsynaptic neuron is at its resting potential, NMDA ion channels are blocked by Mg2+. A model for the mechanism of LTP is illustrated in FIGURE 42-16. A presynaptic neuron releases glutamate, which binds with AMPA receptors. The postsynaptic neuron becomes depolarized. If depolarization of the postsynaptic neuron is sufficiently strong, Mg2+ move away from the NMDA receptors, unblocking them. Glutamate can then bind with these receptors, opening Ca2+ channels and letting Ca2+ move into the cell. Calcium ions appear to be an important trigger for LTP. Calcium ions act as second messengers that initiate long-term changes ultimately responsible for LTP. For example, calcium ions activate a Ca2+-dependent second-messenger pathway that results in the insertion of more AMPA receptors in the postsynaptic membrane. This is important because additional AMPA receptors increase sensitivity to glutamate. More EPSPs (excitatory postsynaptic potentials) are produced, which strengthens the synapse (that is, helps maintain the LTP). Recall from Chapter 41 that an EPSP is a change in membrane potential that brings the neuron closer to its threshold level. Calcium ions also activate a pathway that leads to release of a retrograde signal, a signal that moves backward, from the postsynaptic neuron to the presynaptic neuron. The soluble gas nitric oxide (NO) has been identified as the retrograde signal. This signal enhances neurotransmitter release by the presynaptic neuron. Note that this is a positive feedback loop that strengthens the connection between the two neurons. Long-term memory involves gene expression Gene expression and protein synthesis take place during the process of establishing long-term memory. This process involves slower, but longer-lasting, changes in synaptic connections. Longterm memory depends on activated receptors linked to G proteins (see Chapter 6). Cyclic AMP acts as a second messenger. A high level of cyclic AMP activates a protein kinase that enters the nucleus, leading to gene activation and protein synthesis. In this process, protein kinase phosphorylates a transcription factor known as CREB (for “cyclic AMP response element binding protein”). CREB then turns on the transcription process of certain genes. CREB has been shown to be a signaling molecule in the memory pathway in many animals, including fruit flies and mice. The molecules and processes involved in learning and memory have been highly conserved during evolution. Experience affects development and learning Many studies have demonstrated synaptic plasticity in rats, mice, and other laboratory animals exposed to enriched environments. In contrast with rats housed in standard cages and provided with the basic necessities, those exposed to enriched environments are

KEY POINT

When glutamate binds to NMDA receptors, Ca2+ enter the postsynaptic neuron and activate intracellular signaling systems that lead to changes responsible for LTP. Presynaptic neuron

Presynaptic neuron

Presynaptic neuron

1

Stimulated presynaptic neuron releases glutamate.

Glutamate 4 3

Glutamate Mg 2+

AMPA receptor 2

Glutamate binds with AMPA receptors on postsynaptic neuron.

Postsynaptic neuron

Ion channels open, allowing Na+ to pass into postsynaptic neuron. Na+ depolarize Na+ membrane.

NMDA receptor blocked by Mg 2+

Mg

2+

Ca 2+

If postsynaptic neuron is strongly depolarized, Mg2+ move away from NMDA receptors, allowing glutamate to bind with them.

6

Ca2+ cause more AMPA receptors to be inserted into membrane.

7

Ca 2+

5

Ca2+ channels open.

Signaling pathway NO

(a) Activation of AMPA receptors. Glutamate released by a presynaptic neuron binds with AMPA receptors on the postsynaptic neuron. Note that NMDA receptors are blocked by Mg2+.

(b) Activation of NMDA receptors. The combination of strong stimulation and glutamate binding activates NMDA receptors.

Ca2+ stimulate release of NO, which diffuses into presynaptic neuron and stimulates release of more glutamate.

Retrograde signal

(c) Long-term potentiation. Calcium ions activate pathways, leading to changes that are ultimately responsible for LTP.

FIGURE 42-16 A proposed model for the mechanism of long-term potentiation (LTP)

given toys and other stimulating objects, as well as the opportunity to socially interact with other rats. Animals reared in an enriched environment exhibit increased synaptic contacts and process and remember information more quickly than animals lacking such advantages. Mice exposed to enriched environments develop significantly greater numbers of neurons in the hippocampus and learn mazes faster than control animals. Early environmental stimulation can also enhance the development of motor areas in the brain. For example, the brains of rats encouraged to exercise become slightly heavier than those of control animals. Characteristic changes occur within the cerebellum, including the development of larger dendrites. Apparently, during early life certain critical or sensitive periods of nervous system development occur that are influenced by environmental stimuli. For example, when the eyes of young mice first open, neurons in the visual cortex develop large numbers of dendritic spines (structures on which synaptic contact takes place). If the animals are kept in the dark and deprived of visual stimuli, fewer dendritic spines form. If the mice are exposed to light later in life, some new dendritic spines form but never as many as develop in a mouse reared in a normal environment. Studies linking the development of the human brain with environmental experience have demonstrated that early stimulation is important for the sensory, motor, intellectual, and social development of children. Such studies argue for the importance of informed parenting, as well as for early education programs. Inves-

tigators have also shown that environmental stimulation is needed to maintain the status of the cerebral cortex in later life.

Language involves comprehension and expression Language is a form of communication in which we use words to symbolize objects and convey ideas. In about 97% of humans, including most who are left-handed, the areas of the brain responsible for language are located in the left hemisphere (see Fig. 42-12). Wernicke’s area, located in the temporal lobe, is an important center for language comprehension. This area helps us recognize and interpret both spoken and written words. Wernicke’s area also helps us formulate the choice and sequence of words and transfers information to Broca’s area. Located near motor areas in the left frontal lobe, Broca’s area controls our ability to speak. Broca’s area is also important in language processing and speech comprehension.

Review ■

■ ■ ■

■

How is the human CNS protected? What is the function of the cerebrospinal fluid? What are two main functions of the vertebrate spinal cord? What are the functions of each of the major lobes of the cerebrum? What role does the limbic system play in the emotional aspects of behavior? What role does synaptic plasticity play in information processing?

42.5 THE PERIPHERAL NERVOUS SYSTEM ■ ■ LEARNING OBJECTIVES 10 Describe the organization of the peripheral nervous system and compare its somatic and autonomic divisions.

11 Contrast the sympathetic and parasympathetic divisions of the autonomic system and give examples of the effects of these systems on specific organs such as the heart and intestine.

The peripheral nervous system consists of the sensory receptors, the nerves that link these receptors with the central nervous system, and the nerves that link the CNS with effectors (muscles and glands). The somatic division of the PNS helps the body respond to changes in the external environment. The nerves and receptors that maintain homeostasis despite internal changes make up the autonomic division.

The somatic division helps the body adjust to the external environment The somatic division of the PNS includes the receptors that react to changes in the external environment, the sensory neurons that inform the CNS of those changes, and the motor neurons that adjust the positions of the skeletal muscles that help maintain the body’s posture and balance. In mammals, 12 pairs of cranial nerves emerge from the brain. They transmit information to the brain from the sensory receptors for smell, sight, hearing, and taste. For example, cranial nerve II, the optic nerve, transmits signals from the retina of the eye to the brain. The cranial nerves also transmit information to the brain from general sensory receptors, especially in the head region. Cranial nerves also bring orders from the CNS to the voluntary muscles that control movements of the eyes, face, mouth, tongue, pharynx, and larynx. Cranial nerve VII, the facial nerve, transmits signals to the muscles used in facial expression and to the salivary glands. In humans, 31 pairs of spinal nerves emerge from the spinal cord. Named for the general region of the vertebral column from which they originate, they comprise 8 pairs of cervical, 12 pairs of thoracic, 5 pairs of lumbar, 5 pairs of sacral, and 1 pair of coccygeal spinal nerves. The ventral branches of several spinal nerves form tangled networks called plexi (sing., plexus). Within a plexus, the fibers of a spinal nerve may separate and then regroup with fibers that originated in other spinal nerves. Thus, nerves emerging from a plexus consist of neurons from several different spinal nerves.

to the CNS. The information is integrated at various levels. Then the “decisions” are transmitted along efferent nerves to the appropriate muscles or glands. Afferent and efferent neurons of the autonomic division lie within cranial and spinal nerves. For example, afferent fibers of cranial nerve X, the vagus nerve, transmit signals from many organs of the chest and upper abdomen to the CNS. Its efferent fibers transmit signals from the brain to the heart, stomach, small intestine, and several other organs. The efferent portion of the autonomic division is subdivided into sympathetic and parasympathetic systems (TABLE 42-3). Many organs are innervated by both types of nerves. In general, sympathetic and parasympathetic systems have opposite effects (FIGS. 42-17 and 42-18). For example, the heart rate is speeded up by messages from sympathetic nerve fibers and slowed by impulses from its parasympathetic nerve fibers. In many cases sympathetic nerves operate to stimulate organs and to mobilize energy, especially in response to stress; whereas the parasympathetic nerves influence organs to conserve and restore energy, particularly during quiet, calm activities. Instead of using a single efferent neuron, as in the somatic system, the autonomic system uses a relay of two neurons between the CNS and the effector. The first neuron, called the preganglionic neuron, has a cell body and dendrites within the CNS. Its axon, part of a peripheral nerve, ends by synapsing with a postganglionic neuron. The dendrites and cell body of the postganglionic

KEY POINT

Most organs are innervated by both sympathetic and parasympathetic nerves.

Paravertebral ganglion Vagus nerve (parasympathetic) Speeds Heart Slows Excites Celiac ganglion (part of the solar plexus)

Stomach

Relaxes

The autonomic division regulates the internal environment The autonomic division helps maintain homeostasis in the internal environment. For example, it regulates the heart rate and helps maintain a constant body temperature. The autonomic system works automatically and without voluntary input. Its effectors are smooth muscle, cardiac muscle, and glands. Like the somatic system, it is functionally organized into reflex pathways. Receptors within the viscera relay information via afferent nerves

FIGURE 42-17 Dual innervation of the heart and stomach A sympathetic nerve increases the heart rate, whereas a parasympathetic nerve, the vagus nerve, slows the heartbeat. In contrast, sympathetic nerves slow contractions of the stomach and intestine, whereas the vagus nerve stimulates contractions of the digestive organs. Sympathetic nerves are shown in red; postganglionic fibers are shown as dotted lines.

TABLE 42-3

Comparison of Sympathetic and Parasympathetic Systems Sympathetic System

Parasympathetic System

General effect

Prepares body to cope with stressful situations

Restores body to resting state after stressful situation; actively maintains normal body functions

Extent of effect

Widespread throughout body

Localized

Duration of effect

Lasting

Brief

Outflow from CNS

Thoracic and lumbar nerves from spinal cord

Cranial nerves and sacral nerves from spinal cord

Location of ganglia

Chain and collateral ganglia

Terminal ganglia

Neurotransmitters released

Preganglionic: acetylcholine Postganglionic: norepinephrine (usually)

Acetylcholine Acetylcholine

III VII IX X (vagus nerve)

Medulla

Eyes (pupils) Salivary glands, nasal mucosa, etc.

Cervical

Larynx Lungs Heart

Liver Sweat glands, blood vessels, erector muscles of hair follicles

Gallbladder Adrenal gland

Thoracic

Celiac ganglion

Kidney

Superior mesenteric ganglion

Stomach

Sacral

Lumbar

Intestine Bladder Inferior mesenteric ganglion

Genitals

Sympathetic Parasympathetic Postganglionic nerves Sympathetic ganglia chain

FIGURE 42-18 The sympathetic and parasympathetic systems Complex as it appears, this diagram has been greatly simplified. Red lines represent sympathetic nerves, black lines represent parasympathetic nerves, and dotted lines represent postganglionic nerves.

neuron are in a ganglion outside the CNS. Its axon terminates near or on the effector. The sympathetic ganglia are paired, and a chain of them, the paravertebral sympathetic ganglion chain, runs on each side of the spinal cord from the neck to the abdomen. Some sympathetic preganglionic neurons do not end in these ganglia but pass on to collateral ganglia in the abdomen, close to the aorta and its major branches. Parasympathetic preganglionic neurons synapse with postganglionic neurons in terminal ganglia near or within the walls of the organs they innervate. The sympathetic and parasympathetic systems also differ in the neurotransmitters they release at the synapse with the effector. Both preganglionic and postganglionic parasympathetic neurons secrete acetylcholine. Sympathetic postganglionic neurons release norepinephrine (although preganglionic sympathetic neurons secrete acetylcholine). The autonomic division got its name from the original idea that it was independent of the CNS, that is, autonomous. Neurobiologists have disproved this and shown that the hypothalamus and many other parts of the CNS help regulate the autonomic system. Although the autonomic system usually functions automatically, its activities can be consciously influenced. Biofeedback provides a person with visual or auditory evidence concerning the status of an autonomic body function. For example, a tone may be sounded when blood pressure reaches a desirable level. Using such techniques, individuals have learned to control autonomic activities such as blood pressure, brain wave pattern, heart rate, and blood sugar level. Even certain abnormal heart rhythms can be consciously modified.

tal Health Services Administration, an estimated 20.1 million individuals (8% of the population) ages 12 or older in the United States use drugs that are not legal (based on individuals who acknowledged using an illicit drug at least once during the 30 days prior to the interview). Abuse of prescription drugs, particularly pain killers, has become a serious problem, especially among teens and young adults. Although legal, alcohol is the most commonly used and abused drug. (See Inquiring About: Alcohol: The Most Abused Drug.) More than 9 million children live with a parent who is dependent on alcohol and/or illicit drugs. TABLE 42-4 lists several commonly used and abused drugs and gives their effects. Habitual use of almost any mood-altering drug can result in psychological dependence, in which the user becomes emotionally dependent on the drug. When deprived of it, the user craves the feeling of euphoria (well-being) the drug induces. Some drugs induce tolerance after several days or weeks. Response to the drug decreases, and greater amounts are required to obtain the desired effect. Tolerance often occurs because the liver cells are stimulated to produce more of the enzymes that metabolize and inactivate the drug. It can also occur when the number of postsynaptic receptors that bind with the drug decrease (down-regulation). Drug addiction is a serious societal problem that involves compulsive use of a drug despite negative health consequences and negative effects on the ability to function socially and occupationally. Use of some drugs, such as heroin, tobacco, alcohol, and barbiturates, may also result in physical dependence, in which physiological changes occur that make the user dependent on the drug. For example, when heroin or alcohol is withheld, the addict suffers characteristic withdrawal symptoms. Physical addiction can occur when a drug, for example, morphine, has components similar to substances that body cells normally manufacture on their own. Some highly addictive drugs, such as crack cocaine and methamphetamine, do not cause serious withdrawal symptoms.

Review ■

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What are some differences between the somatic division and the autonomic division? What are some differences in structure and function of the sympathetic system and the parasympathetic system?

42.6 EFFECTS OF DRUGS ON THE NERVOUS SYSTEM ■ ■ LEARNING OBJECTIVE

Neural signal Presynaptic neuron

Dopamine reuptake

Dopamine released and binds to receptors.

12 Distinguish among psychological dependence, tolerance, and drug addiction, and describe the biological actions and effects on mood of alcohol, antidepressants, barbiturates, antianxiety drugs, antipsychotic drugs, opiates, stimulants, hallucinogens, and marijuana.

About 25% of all prescribed drugs are taken to alter psychological conditions, and almost all the commonly abused drugs affect mood. Many act by changing the levels of neurotransmitters within the brain. In particular, changing levels of norepinephrine, serotonin, and dopamine influence mood. According to the Substance Abuse and Men-

Neural signal Presynaptic neuron

Cocaine blocks reuptake.

Dopamine accumulates and binds to receptors. More neural signals

Neural signal

Postsynaptic neuron

(a) Reuptake of the neurotransmitter dopamine. Dopamine binds with a transporter (green) that shuttles it back into the presynaptic terminal.

FIGURE 42-19 Mechanism of cocaine action

Postsynaptic neuron

(b) Cocaine blocks the reuptake of dopamine by binding competitively with the dopamine reuptake transporter. As a result, more dopamine is available to bind with postsynaptic receptors.

In q u ir in g A b o u t A LCO H O L : T H E M OS T A B US E D D RU G What makes alcohol one of the leading preventable risk factors for premature death in the United States? An estimated 90,000 individuals in the United States alone die each year as a result of alcohol-related cancers and other diseases, alcohol-related violence, traffic accidents, and other injuries. Alcohol abuse costs our society more than $100 billion annually. An estimated 18 million adults in the United States have problems with alcohol abuse or dependence. More than half of the adults in the United States report that one or more of their close relatives has a drinking problem. According to the most recent U.S. Department of Health and Human Services study, the 2008 National Survey on Drug Use and Health: National Findings, 23% of Americans had binged (five or more drinks within two hours) at least once during the 30 days prior to being surveyed. More than 23% had drunk heavily (five or more drinks on five or more occasions). The highest prevalence of binge and heavy drinking was among young adults ages 18 to 25 years. More than seven million binge drinkers in the United States are under age 21. According to the U.S. Department of Health and Human Services, alcohol use by individuals under age 21 is a major public health problem. Adolescents ages 12 to 20 consume 11% of all alcohol consumed in the United States, and more than 90% of this alcohol was consumed during bingeing. Based on many studies, the Centers for Disease Control and Prevention (CDC) reports that youth who drink alcohol are likely to experience serious problems, including disruption of normal growth and sexual development, abnormal brain development that may have life-long effects, memory problems, as well as school, social, and legal problems and higher risk for suicide, homicide, and death from alcohol poisoning. Investigators have reported that adults who first used alcohol when they were 14 years old or younger were more than five times as likely to have a problem with

alcohol dependence or abuse than adults who had their first drink at age 21 or older. Alcohol abuse results in physiological, psychological, and social impairment and has serious negative consequences for family, friends, and society. Alcohol abuse has been linked to the following: • More than 30% of all traffic fatalities • Almost 50% of violent crime • More than 60% of child-abuse and spouse-abuse cases • More than 50% of college sexual assaults • About 25% of suicides • The birth each year of thousands of babies with serious birth defects because their mother drank alcohol during pregnancy. Fetal alcohol spectrum disorders are the leading cause of preventable mental retardation in the United States. • Increased risk of mouth, larynx, and throat cancers. Alcohol is also a risk factor for primary liver cancer, breast cancer, and colon cancer. These cancers have been linked to regular alcohol consumption. A single drink, that is, 12 oz of beer, 5 oz of wine, or 1.5 oz of 80-proof liquor, results in a blood alcohol concentration of approximately 20 mg/dL (milligrams per deciliter). This represents about 0.5 oz of pure alcohol in the blood. Alcohol accumulates in the blood because absorption occurs more rapidly than oxidation and excretion. Every cell in the body can take in alcohol from the blood. At first the drinker may feel stimulated, but alcohol actually depresses the CNS. As blood alcohol concentration rises, information processing, judgment, memory, sensory perception, and motor coordination all become progressively impaired. Depression and drowsiness generally occur. Contrary to popular belief, alcohol decreases sexual performance. Some individuals become loud, angry, or violent.

The neurobiological mechanisms for drug addiction involve the mesolimbic dopamine pathway, a major reward circuit (described earlier in this chapter in connection with the limbic system). Many of the neurons in this pathway project from the ventral tegmental area in the midbrain and signal neurons in the limbic system, including the amygdala and an area called the nucleus accumbens. Projections of this nucleus are thought to be involved in the reinforcement people experience from taking drugs. Effects on neurons in the mesolimbic dopamine pathway have been demonstrated in nicotine, heroin, amphetamine, and cocaine addiction. For example, amphetamines stimulate dopamine

A blood alcohol concentration of 80 mg/ dL (or 0.08 gm/100 mL) legally defines driving under the influence (DUI). A 170-pound man typically reaches this level by drinking four drinks in an hour on an empty stomach. Alcohol metabolism occurs at a fixed rate in the liver so only time, not coffee, decreases its effects. Because alcohol inhibits water reabsorption in the kidneys, more fluid is excreted as urine than is consumed in the alcohol. This results in dehydration that, together with low blood sugar level, may cause the stupor produced by excessive drinking. In chronic drinkers, cells of the CNS adapt to the presence of the drug. This causes tolerance (more and more alcohol is needed to experience the same effect) and physical dependence. Abrupt withdrawal can result in sleep disturbances, severe anxiety, tremors, seizures, hallucinations, and psychoses. The A1 allele of the D2 dopamine receptor gene increases the risk for alcoholism. Individuals with this allele have fewer D2 dopamine receptors in their brain. Alcohol, like cocaine, heroin, amphetamines, and nicotine, enhances dopamine activity in the mesolimbic dopamine pathway of the brain. Individuals with the A1 allele may compensate for their low dopamine levels by using alcohol and other drugs. However, the mechanisms underlying alcohol addiction are far more complex. Evidence suggests that many neurotransmitters are involved, including glutamate, serotonin, GABA, and opioid peptides. Recently, researchers identified a cluster of genes on chromosome 11 that appear to be associated with alcohol dependence. Treatment for alcohol problems includes various forms of psychotherapy and use of drugs such as naltrexone and acamprosate. The group support offered by Alcoholics Anonymous (AA) has proved effective for many struggling with alcohol dependence. Participation in AA includes attending frequent meetings, having a sponsor within the program, and working on the 12 steps.

release. Cocaine blocks the reuptake of dopamine into presynaptic neurons, thus prolonging the stimulation (FIG. 42-19). Amphetamines or cocaine can increase dopamine concentration in the limbic system by up to 1000 times normal amounts! Prolonged drug use alters gene expression and changes neuron structure.

Review ■

How does each of the following drugs affect the CNS: (1) alcohol, (2) antipsychotic drugs, (3) antidepressants, (4) amphetamines, and (5) MDMA (“Ecstasy”)?

TABLE 42-4

Effects of Some Commonly Used Drugs

Name of Drug

Side Effects/Dangers Associated with Abuse

Effect on Mood

Actions on Body

Tricyclic antidepressants (e.g., Elavil, Anafranil)

Elevate mood; relieve depression; used to treat obsessivecompulsive behavior

Most block reuptake of biogenic amines (especially norepinephrine), increasing their concentration in synapses

Sedation, weight gain, sexual dysfunction

Selective serotonin reuptake inhibitors (SSRIs) (e.g., Prozac, Zoloft, Lexapro, Celexa)

Relieve depression; used to treat obsessive-compulsive behavior

Block serotonin reuptake

Nausea, headache, insomnia, anxiety

MAO (monoamine oxidase) inhibitors (e.g., Parnate, Nardil)

Relieve depression

Block enzymatic breakdown of biogenic amines, increasing their concentration in synapses

Liver toxicity, excessive CNS stimulation; overdose may affect blood pressure, cause hallucinations

Benzodiazepines (e.g., Xanax, Valium, Librium)

Sedation; induce sleep

Bind to GABA receptors on postsynaptic neuron; increase effectiveness of GABA in opening chloride channels, causing hyperpolarization; cause relaxation of skeletal muscles

Drowsiness, confusion; psychological dependence, addiction; effects additive with alcohol

Barbiturates (“downers”) (e.g., phenobarbital)

Sedation; induce sleep

Bind to receptor adjacent to GABA receptor on chloride channels; chloride channels open so more chloride ions move in, causing hyperpolarization

Drowsiness, confusion; psychological dependence, tolerance, addiction; severe CNS depression, resulting in coma and death (especially lethal in combination with alcohol)

Phenothiazines (e.g., Thorazine, Mellaril, Stelazine)

Relieve symptoms of schizophrenia; reduce impulsive and aggressive behavior

Block dopamine receptors

Muscle spasms, shuffling gait, and other symptoms of Parkinson’s disease

Atypical antipsychotic medications (e.g., Abilify, Risperdal)

Similar to phenothiazines, but also increase motivation

Block dopamine receptors and other neurotransmitter receptors

Fewer side effects than with the phenothiazines

Euphoria; sedation; relieve pain

Mimic actions of endorphins; bind to opiate receptors

Depress respiration; constrict pupils; impair coordination; tolerance, psychological dependence, addiction; convulsions, death from overdose

Cocaine

Brief, intense high with euphoria, followed by fatigue and depression

Stimulates release and inhibits reuptake of norepinephrine and dopamine, leading to CNS stimulation followed by depression; autonomic stimulation; dilation of pupils; local anesthesia

Mental impairment, convulsions, hallucinations, unconsciousness; death from overdose

Marijuana

Euphoria

Impairs coordination; impairs depth perception and alters sense of timing; impairs short-term memory (probably by decreasing acetylcholine levels in the hippocampus); inflames eyes; causes peripheral vasodilation

In large doses, sensory distortions, hallucinations, evidence of lowered sperm counts and testosterone levels

Euphoria, stimulation, hyperactivity

Stimulate release and block reuptake of dopamine and norepinephrine; enhance flow of impulses in RAS, leading to increased heart rate, blood pressure; pupil dilation

Tolerance, physical dependence; hallucinations; increased blood pressure; psychotic episodes; death from overdose

Antidepressants

Anti-anxiety Medications

Antipsychotic Medications

Narcotic Analgesics Opiates (e.g., morphine, codeine, Demerol, heroin)

Amphetamines “Uppers,” “pep pills,” “crystal meth” (e.g., Dexedrine); methamphetamine is a popular club drug

TABLE 42-4

(Continued)

Name of Drug

Side Effects/Dangers Associated with Abuse

Effect on Mood

Actions on Body

Amphetamine-type prescription stimulants commonly used to treat attention deficit hyperactivity disorder (ADHD) (e.g., methylphenidate: Ritalin, Concerta)

CNS stimulation, but has calming effect on many children and adults who have ADHD; increases focus

Increases heart rate, blood pressure, and body temperature; like cocaine, amphetamines block dopamine transporter which increases dopamine concentration in synapses

Not addictive at prescribed doses; used illegally and abused by some high school and college students who use them in higher doses than would be prescribed; can cause irregular heartbeat and respiration, hallucinations, delusions, and convulsions; abuse can lead to tolerance and physical addiction

MDMA (3,4-methylenedioxymethamphetamine) “Ecstasy” (most popular of club drugs)

Stimulation, sense of pleasure, self-confidence, feelings of closeness to others; mild hallucinogenic

CNS stimulant; damages serotonin pathways in rat and monkey brains

Hyperthermia (excessive body heat); overdose causes rapid heartbeat, high blood pressure, panic attacks, and seizures; longterm neurological changes; cognitive impairment

Rohypnol (flunitrazepam) “Date rape” drug, “roofies,” “roach”

Sedative-hypnotic

Muscle relaxation; amnesia

Physical and psychological dependence; withdrawal may cause seizures; may be lethal when mixed with alcohol and/or other depressants

PCP (phencyclidine) “Angel dust,” “ozone”

Feelings of strength, power, invulnerability, and a numbing effect on the mind

Increase in blood pressure and heart rate; shallow respiration; flushing, profuse sweating; numbness of extremities; muscular incoordination; changes in body awareness, similar to those associated with alcohol intoxication

Psychological dependence, craving; in adolescents, interferes with hormones related to growth and development; may interfere with learning; causes hallucinations, delusions, disordered thinking; high doses cause drop in blood pressure and heart rate, seizures, coma, and death; chronic use causes memory loss, difficulties with speech and thinking, and depression

LSD (lysergic acid diethylamide)

Overexcitation; sensory distortions, hallucinations

Alters levels of neurotransmitters in brain; potent CNS stimulator; dilates pupils, sometimes unequally; increases heart rate; raises blood pressure

Impaired judgment leading to irrational behavior; panic attacks; psychosis; flashbacks

Methaqualone (e.g., Quaalude, Sopor)

Hypnotic

Depresses CNS; depresses certain spinal reflexes

Tolerance, physical dependence; convulsions, death

Caffeine

Increases mental alertness; decreases fatigue and drowsiness

Acts on cerebral cortex; relaxes smooth muscle; stimulates cardiac and skeletal muscle; increases urine volume

Very large doses stimulate centers in the medulla (may slow the heart); toxic doses may cause convulsions

Nicotine

Decreases psychological tension

Stimulates sympathetic nervous system; increases dopamine in mesolimbic pathway

Tolerance, physical dependence; stimulates development of atherosclerosis by stimulating synthesis of lipid in arterial wall

Alcohol

Euphoria, relaxation, release of inhibitions

Depresses CNS; impairs vision, coordination, judgment; lengthens reaction time

Physical dependence; damage to pancreas and liver (e.g., cirrhosis); brain damage; increased risk of depression

Amphetamines (continued )

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

42.1 (page 883)

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1 Contrast nerve nets and radial nervous systems with bilateral nervous systems. Nerve nets and radial nervous systems are typical of radially symmetrical invertebrates, such as cnidarians. An animal with a radial nervous system can respond effectively to predator or prey approaching from any direction. ■ In planarian flatworms the bilateral nervous system includes cerebral ganglia and, usually, two solid ventral nerve cords connected by transverse nerves. Annelids and arthropods typically have a ventral nerve cord and numerous ganglia. The cerebral ganglia of arthropods have specialized regions. Octopods and other cephalopod mollusks have highly developed nervous systems with neurons concentrated in a central region. ■ The bilateral nervous system is adaptive for bilaterally symmetrical animals that typically move in a forward direction. Nerve cells concentrate to form nerves, nerve cords, ganglia, and brain; typically, sense organs are concentrated in the head region. ■

Learn more about various nervous systems by clicking on the figures in CengageNOW.

2 Identify trends in the evolution of the invertebrate nervous system. ■

Trends in nervous system evolution include increased numbers and concentration of nerve cells; specialization of function; increased number of association neurons; more complex synaptic contacts; and cephalization, formation of a head.

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Explore the early development of the vertebrate nervous system and the evolution of the vertebrate brain by clicking on the figures in CengageNOW.

42.4 (page 888) 5 Describe the structure and functions of the human spinal cord. ■

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42.2 (page 884) 3 Describe the two main divisions of the vertebrate nervous system and summarize their functions. The central nervous system (CNS), which consists of the brain and spinal cord, is specialized to integrate incoming information. The peripheral nervous system (PNS), which consists of sensory receptors and nerves, transmits signals between sensory receptors and the CNS, and between the CNS and effectors.

The human brain and spinal cord are protected by bone and three meninges—the dura mater, arachnoid, and pia mater; the brain and spinal cord are cushioned by cerebrospinal fluid (CSF). The spinal cord transmits impulses to and from the brain and controls many reflex actions. The spinal cord consists of ascending tracts, which transmit information to the brain, and descending tracts, which transmit information from the brain. Its gray matter contains nuclei that serve as reflex centers. Learn more about the spinal cord by clicking on the figure in CengageNOW.

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Learn more about the vertebrate nervous system by clicking on the figure in CengageNOW.

The forebrain differentiates to form the diencephalon and telencephalon. The diencephalon develops into the thalamus and hypothalamus. The thalamus is a relay center for motor and sensory information. The hypothalamus controls autonomic functions; links nervous and endocrine systems; controls temperature, appetite, and fluid balance; and is involved in some emotional and sexual responses. The telencephalon develops into the cerebrum and olfactory bulbs. In most vertebrates, the cerebrum is divided into right and left cerebral hemispheres. In fishes and amphibians, the cerebrum mainly integrates incoming sensory information. In birds, the corpus striatum controls complex behavior patterns, such as flying and singing. In mammals, the neocortex accounts for a large part of the cerebral cortex, the gray matter of the brain. The cerebrum has complex association functions.

A withdrawal reflex involves sensory receptors; sensory neurons, interneurons, and motor neurons; and effectors, such as muscles. 6 Describe the structure and functions of the human cerebrum. ■

Parietal lobe

42.3 (page 886)

Central sulcus

Frontal lobe

4 Trace the development of the principal vertebrate brain regions from the hindbrain, midbrain, and forebrain, and compare the brains of fishes, amphibians, reptiles, birds, and mammals. In the vertebrate embryo, the brain and spinal cord arise from the ■ neural tube. The anterior end of the tube differentiates into hindbrain, midbrain, and forebrain. ■ The hindbrain subdivides into the metencephalon and myelencephalon. The myelencephalon develops into the medulla, which contains vital centers and other reflex centers. The fourth ventricle, the cavity of the medulla, communicates with the central canal of the spinal cord. The metencephalon gives rise to the cerebellum, which is responsible for muscle tone, posture, and equilibrium, and to the pons, which connects various parts of the brain. ■ The midbrain is the largest part of the brain in fishes and amphibians. It is their main association area, linking sensory input and motor output. In reptiles, birds, and mammals, the midbrain serves as a center for visual and auditory reflexes. The medulla, pons, and midbrain make up the brain stem.

Prefrontal area Occipital lobe Cerebellum Temporal lobe Brain stem Medulla

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The human cerebral cortex consists of gray matter, which forms folds or convolutions. Deep furrows between the folds are called fissures. The cerebrum is functionally divided into sensory areas that receive incoming sensory information; motor areas that control voluntary movement; and association areas that link sensory and motor areas and are responsible for learning, language, thought, and judgment. The cerebrum consists of lobes, including the frontal lobes, parietal lobes, temporal lobes, and occipital lobes. The white matter of the cerebrum lies beneath the cerebral cortex. The corpus callosum, a large band of white matter, connects right and left hemispheres. The basal ganglia, a cluster of nuclei within the white matter, are important centers for motor function. Learn more about the human brain by clicking on the figures in CengageNOW.

7 Summarize the sleep–wake cycle and contrast rapid-eye-movement (REM) sleep with non-REM sleep. ■ The body’s main biological clock—the suprachiasmatic nucleus— receives information about light and dark and transmits it to other nuclei that regulate sleep. ■ The reticular activating system (RAS) is an arousal system; its neurons filter sensory input, selecting which information is transmitted to the cerebrum. ■ The hypothalamus, thalamus, and brain stem regulate the sleep– wake cycle. Sleep is an alteration of consciousness with decreased electrical activity in the cerebral cortex. ■ Electrical activity of the cerebral cortex and metabolic rate slow during non-REM sleep, whereas dreaming characterizes REM sleep. 8 Describe the actions of the limbic system, including the role of the amygdala in emotional expression. ■ The limbic system affects the emotional aspects of behavior, motivation, sexual activity, autonomic responses, and biological rhythms. The amygdala evaluates incoming information and signals danger; it is especially important in the experience of fear and aggression. 9 Summarize how the brain processes information, including a description of synaptic plasticity and the neurobiological changes that take place during learning. ■ Synaptic plasticity is the ability of the nervous system to modify synapses, which allows learning and remembering. ■ Learning is the process by which we acquire knowledge or skills as a result of experience. Memory is the process by which information is encoded, stored, and retrieved. Implicit memory is unconscious

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(“how-to”) memory for perceptual and motor skills. Declarative memory is factual memory of people, places, or objects. Immediate memory holds sensory information for fractions of a second. Short-term memory holds a small amount of information, for a few minutes. Information can be transferred from short-term memory to long-term memory. The hippocampus is important in categorizing information and consolidating memories. Learning and memory involve long-term functional changes at synapses. Known as long-term potentiation (LTP), such changes involve increased sensitivity to an action potential by a postsynaptic neuron. Long-term depression (LTD) is a long-lasting decrease in the strength of synaptic connections. Long-term memory storage involves gene expression. The transcription factor CREB is an important signaling molecule in the memory pathway.

42.5 (page 902) 10 Describe the organization of the peripheral nervous system and compare its somatic and autonomic divisions. The PNS consists of sensory receptors and nerves, including the cranial nerves and spinal nerves and their branches. The somatic division of the PNS responds to changes in the external environment. The autonomic division regulates the internal activities of the body. 11 Contrast the sympathetic and parasympathetic divisions of the autonomic system and give examples of the effects of these systems on specific organs such as the heart and intestine. ■ The sympathetic system permits the body to respond to stressful situations. The parasympathetic system influences organs to conserve and restore energy. Many organs are innervated by sympathetic and parasympathetic nerves, which function in opposite ways. For example, the sympathetic system increases heart rate, whereas the parasympathetic system decreases heart rate. ■

42.6 (page 904) 12 Distinguish among psychological dependence, tolerance, and drug addiction, and describe the biological actions and effects on mood of alcohol, antidepressants, barbiturates, anti-anxiety drugs, antipsychotic drugs, opiates, stimulants, hallucinogens, and marijuana. ■ Many drugs alter mood by increasing or decreasing the concentrations of specific neurotransmitters within the brain. See Table 42-4 for a summary of these drugs. ■ Habitual use of mood-altering drugs can result in psychological dependence or drug addiction. Many drugs induce tolerance, in which the body’s response to the drug decreases so that greater amounts are needed to obtain the desired effect.

T E S T YO U R U N D E R S TA N D I N G 1. A radially symmetrical animal such as Hydra is likely to have (a) a forebrain (b) a nerve net (c) cerebral ganglia (d) a ventral nerve cord (e) cerebral ganglia and a nerve net

5. Which part of the brain controls autonomic functions and regulates body temperature? (a) cerebrum (b) hypothalamus (c) cerebellum (d) pons (e) thalamus

2. In vertebrate embryos the brain develops from the (a) spinal cord (b) sympathetic nervous system (c) parasympathetic nervous system (d) neural tube (e) forebrain 3. Which part of the brain maintains posture, muscle tone, and equilibrium? (a) cerebrum (b) medulla (c) cerebellum (d) neocortex (e) thalamus

6. Which sequence best describes transmission of information in a withdrawal reflex? 1. afferent neuron transmits signal to CNS 2. muscle contracts 3. reception by sensory neuron 4. interneuron in CNS integrates information 5. efferent neuron transmits signal (a) 3, 1, 4, 5, 2 (b) 2, 1, 4, 5, 3 (c) 3, 5, 1, 4, 2 (d) 3, 1, 5, 4, 2 (e) 5, 3, 1, 4, 2

4. The most prominent part of the amphibian brain is the (a) midbrain (b) medulla (c) cerebellum (d) neocortex (e) cerebrum

7. The human brain is protected by (a) arachnoid, pia mater, cerebrospinal fluid, and ganglia (b) meninges and skull bones only

(c) dura mater and fourth ventricle (d) pia mater and skull bones (e) meninges, cerebrospinal fluid, and skull bones

12. Label the diagram. Use Figure 42-11a to check your answers.

8. The suprachiasmatic nucleus (a) secretes melatonin (b) stimulates beta waves (c) filters incoming sensory information and determines whether it is threatening (d) is critical in memory consolidation (e) is important in regulating the sleep–wake cycle 9. Long-term potentiation (LTP) (a) is mainly the responsibility of cranial nerve II (b) is a long-lasting increase in synaptic strength (c) is associated with short-term memory (d) is a long-lasting decrease in synaptic strength (e) is a long-lasting decrease in synaptic strength that occurs mainly during REM sleep 10. The heart rate is slowed by (a) sympathetic nerves (b) parasympathetic nerves (c) rapid firing in the corpus callosum (d) both sympathetic and parasympathetic nerves (e) the hippocampus working together with sympathetic nerves 11. Amphetamines (a) affect neurons in the mesolimbic dopamine pathway (b) decrease the amount of dopamine secreted (c) have the same types of effects as alcohol (d) are often prescribed to reduce anxiety (e) are CNS depressants

CRITICAL THINKING 1. What is the adaptive function of the limbic system when you are in danger? 2. Hypothesize a possible relationship between inheritance and intelligence based on gene activation during information processing. 3. EVOLUTION LINK. What general trends can you identify in the evolution of the vertebrate brain? 4. EVOLUTION LINK. CREB has been shown to be a signaling molecule in the memory pathway in many animals, including fruit

flies and mice. What does this suggest about the evolution of learning and memory? 5. SCIENCE, TECHNOLOGY, AND SOCIETY. Imagine that you have just become a parent. What kinds of things can you do to enhance the development of your child’s intellectual abilities? Additional questions are available in CengageNOW at www.cengage.com/ login.

Sensory Systems

Merlin D. Tuttle/Bat Conservation International/Photo Researchers, Inc.

43

KEY CONCEPTS

Bats are guided by echoes. This California leaf-nosed bat (Macrotus californicus) has located an insect.

H

ave you heard dolphins make clicking sounds? Humans can hear only a limited number of the wide range of sound frequencies that

43.1 Sensory processing includes sensory reception,

dolphins emit. Dolphins, bats, and a few other vertebrates detect distant

energy transduction, transmission of signals, and interpretation in the central nervous system.

objects by echolocation, sometimes called biosonar. Echolocation works

43.2 Thermoreceptors respond to heat and cold.

ample, a bat emits high-pitched sounds that bounce off objects in its path

43.3 Some predatory species of sharks, rays, and bony

and echo back. By rapidly responding to the echo, the bat skillfully avoids

fishes have electroreceptors that detect electric fields generated in the water by the activity of other animals. Electromagnetic receptors are electroreceptors that detect Earth’s magnetic field.

something like radar but uses sound rather than radio waves. For ex-

obstacles and captures prey (see photograph). Using echolocation, bats and dolphins determine the size, shape, texture, and density of an object, as well as its location. The ability to detect sounds emitted by some foraging bats has

43.4 Dendrites of certain sensory neurons are nociceptors (pain receptors).

evolved in certain moths and other insect prey. Through the course of

43.5 Mechanoreceptors respond to touch, pressure, grav-

evolution, some species of bats have adapted to this ability of their prey;

ity, stretch, or movement. Most animals have mechanoreceptors that detect motion and position; in vertebrates, the vestibular apparatus in the inner ear helps maintain equilibrium, and the organ of Corti within the cochlea of the inner ear contains auditory receptors.

they now emit even higher-frequency sounds that their prey cannot

43.6 Most animals have chemoreceptors for taste and smell

detect. Researchers have recently reported another interesting adaptation. There is evidence that some moths emit ultrasonic clicks that jam the bats’ echolocation. Sensory systems transfer information from the outside world to the

that detect chemical substances in food, water, and air.

animal. They inform the animal about predators, food, water, possible

43.7 Almost all animals have photoreceptors that respond

mates, temperature, and other changes in the environment. The kinds of

to light, and most animals have eyes that form images; in vertebrates, the retina of the eye contains the photoreceptor cells, the rods and cones.

sensory receptors an animal has and the way in which the brain perceives incoming sensory information determine what the animal’s world is like.

We humans live in a world of rich colors, numerous shapes, and varied sounds. But we cannot hear the high-pitched whistles that are audible to dogs and cats or the ultrasonic echoes by which bats navigate. Nor do we ordinarily recognize our friends by their distinctive odors! And although vision is our dominant and most refined sense, we are blind to the ultraviolet (UV) hues that light up the world for insects. Unlike birds and some other animals, we cannot detect changes in magnetic fields. Nor can we detect electric fields like sharks and several other species of fishes. We humans are most familiar with the senses of sight, hearing, smell, taste, and touch, but there are other senses. In fact, touch is a compound sense that allows us to detect pressure, pain, and temperature. Some sensory receptors allow us to sense balance, muscle tension, and joint position, providing us with an awareness of our bodies. In this chapter we will first examine how sensory systems function. Next we will introduce the major types of sensory receptors. We then discuss several types of sensory receptors with emphasis on auditory and visual receptors.

43.1 HOW SENSORY SYSTEMS WORK ■ ■ LEARNING OBJECTIVES 1

2

Describe how sensory systems function; include descriptions of sensory receptors, energy transduction, receptor potential, sensory adaptation, and perception. Classify sensory receptors according to the location of stimuli to which they respond and according to the types of energy they transduce.

Sensory receptors detect information about changes in the external and internal environment. These receptors consist of specialized neuron endings or specialized cells in close contact with neurons. Sensory receptors, along with other types of cells, make up complex sense organs, such as eyes, ears, nose, and taste buds. A human taste bud, for example, consists of modified epithelial cells that detect chemicals dissolved in saliva. Several steps take place in sensory processing, including sensory reception, energy transduction, transmission of the signal, and interpretation in the brain. With minor variations, this is how all sensory systems operate.

Sensory receptors receive information Sensory receptors receive stimuli from the external or internal environment. In the process of reception, they absorb a small amount of energy from some stimulus. Each kind of sensory receptor is especially sensitive to one particular form of energy. The photoreceptors of the human eye are stimulated by an extremely faint beam of light, and taste receptors are stimulated by a minute amount of a chemical compound.

Sensory receptors transduce energy Sensory receptors transduce, or convert, the energy of the stimulus into electrical signals, the information currency of the nervous system. This process is known as energy transduction. When unstimulated, a sensory receptor maintains a resting potential, that is, a difference in charge between the inside and outside of the cell (see Chapter 41). Transduction couples a stimulus with the opening or closing of ion channels in the plasma membrane of sensory receptors. The permeability of the plasma membrane to various ions is altered. A change in ion distribution causes a change in voltage across the membrane. If the difference in charge increases, the receptor becomes hyperpolarized. If the potential decreases, the receptor becomes depolarized. A change in membrane potential is a receptor potential. A receptor potential does not directly trigger an action potential. Like an excitatory postsynaptic potential, or EPSP (see Chapter 41), the receptor potential is a graded response in which the magnitude of change depends on the energy of the stimulus. If the receptor is a separate cell, receptor potentials stimulate the release of a neurotransmitter, which flows across the synapse and binds to receptors on a sensory neuron, also known as an afferent neuron (FIG. 43-1). When a sensory neuron becomes sufficiently depolarized to reach its threshold level, an action potential is generated. Thus, receptor potentials can generate action potentials that transmit information to the central nervous system (CNS). Many receptors are specialized neurons rather than separate cells. In these neurons, the specialized region of the plasma membrane that transduces energy does not generate action potentials. Current generated by receptor potentials flows to a region along the axon where an action potential can be generated. We can summarize the process of sensory reception as follows: stimulus (such as light energy) ¡ sensory receptor absorbs energy from stimulus ¡ transduces energy of stimulus into electrical energy ¡ receptor potential ¡ action potential in sensory neuron ¡ signal transmitted to CNS

Sensory input is integrated at many levels Sensory integration begins in the receptor itself. For example, receptor potentials produced by various stimuli are integrated by summation (see Chapter 41). However, integration occurs mainly in the spinal cord and at various locations in the brain. Sensory receptors adapt to stimuli Many sensory receptors do not continue to respond at the initial rate, even if the stimulus continues at the same intensity. This change in response rate, called sensory adaptation, occurs for two reasons. First, during a sustained stimulus, the receptor sensitivity decreases and produces a smaller receptor potential (resulting in a lower frequency of action potentials in the sensory neurons). Second, changes take place at synapses in the neural pathway activated by the receptor. For example, the release of neurotransmitter from

Stimulus

Sugar molecules

2

Reception. Taste receptor receives stimuli

Receptor potential

Taste receptor

Membrane potential (mV)

1

Energy transduction; receptor potentials generated

Neurotransmitter

1

2

3 4 5 Time (msec)

6

0

Energy transduction; receptor potentials generated

3

Transmission. Sensory neuron depolarized; action potentials generated; signals transmitted to CNS

−60

0

4

2

7

Action potentials

Transmission. Sensory neuron depolarized; action potentials generated; signals transmitted to CNS

Reception. Dendrites in skin receive pain stimuli

−60

0

Membrane potential (mV)

3

1

−50

1

2

3 4 5 Time (msec)

6

7

Integration in brain, leading to perception of taste 4

(a) Taste reception

Integration in spinal cord and brain, leading to perception of pain (b) Pain reception

CNS

FIGURE 43-1 How sensory systems work Sensory receptors receive stimuli from the environment. They transduce the energy of the stimulus into electrical signals, and thus generate receptor potentials. Action potentials are generated in sensory neurons that transmit signals to the CNS, where they are integrated.

a presynaptic terminal may decrease in response to a series of action potentials. Some receptors, such as those for pain or cold, adapt so slowly that they continue to trigger action potentials as long as the stimulus persists. Other receptors adapt rapidly, permitting us to ignore persistent unpleasant or unimportant stimuli. For example, when you first pull on a pair of tight jeans, your pressure receptors let you

know that you are being squished, and you may feel uncomfortable. Soon, though, these receptors adapt, and you hardly notice the sensation of the tight fit. In the same way, people quickly adapt to odors that at first seem to assault their senses. Sensory adaptation enables an animal to discriminate between unimportant background stimuli that can be ignored and new or important stimuli that require attention.

Sensation depends on transmission of a coded message How do you know whether you are seeing a blue sky, tasting a doughnut, or hearing a note played on a piano? All action potentials are qualitatively the same. Light of the wavelength 450 nm (blue), sugar molecules (sweet), and sound waves of 440 hertz (Hz) (A above middle C) all cause transmission of similar action potentials. Our ability to differentiate stimuli depends on both the sensory receptor itself and on the brain. We can distinguish the color of a blue sky from the scent of cologne; a sweet taste from a light breeze; or the sound of a piano from the heat of the sun because cells of each sensory receptor are connected to specific neurons in particular parts of the brain. Because a receptor normally responds to only one type of stimulus (for example, light), the brain interprets a message arriving from a particular receptor as meaning that a certain type of stimulus occurred (such as a flash of color). When stimulated, a sensory receptor initiates what might be considered a coded message, composed of action potentials transmitted by sensory neurons. This coded message is later decoded in the brain. Impulses from the sensory receptor may differ in the (1) total number of sensory neurons transmitting the signal, (2) specific neurons transmitting action potentials and their targets, (3) total number of action potentials transmitted by a given neuron, and (4) frequency of the action potentials transmitted by a given fiber. The intensity of a stimulus is coded by the frequency of action potentials fired by sensory neurons during the stimulus. A strong stimulus results in a greater depolarization of the receptor membrane, so the sensory neuron fires action potentials with greater frequency than would a weak stimulus. This variation is possible because, as a graded response, the receptor potential can vary in magnitude. The difference in sound intensity between the gentle rustling of leaves and a clap of thunder depends on the number of neurons transmitting action potentials as well as the frequency of the action potentials transmitted by each neuron. Just how the sensory receptor initiates different codes and how the brain analyzes and interprets them to produce various sensations are not yet completely understood. Sensation takes place in the brain. Interpretation of the message and the type of sensation depends on which interneurons receive the message. The rods and cones of the eye do not see. When stimulated, they send a message to the brain that interprets the signals and translates them into a rainbow, an elephant, or a child. Artificial (for example, electrical) stimulation of brain centers can also result in sensation. In contrast, many sensory messages never give rise to sensations at all. For example, certain chemoreceptors sense internal changes in the body but never stir our consciousness. Sensory perceptions are constructed by the mind The brain interprets sensations by converting them to perceptions of stimuli. Sensory perception is the process of selecting, interpreting, and organizing sensory information. Sensory perceptions are constructed by the mind by comparing present sensory experience with our memories of past experiences. You may perceive the incoming visual and auditory input of a hissing snake very differently than does your friend who is terrified of snakes, or a herpetologist who studies them. Perceptions are influenced by our state of mind at the time we receive sensory information. Optical illu-

(a) Your perception of the same visual input can vary. Do you see a wineglass or the profiles of two faces?

(b) Do you “see” a white square? If so, your brain is completing a familiar pattern suggested by the right-angle wedges taken out of each of the four red circles.

FIGURE 43-2 Visual perception

sions demonstrate that the brain can interpret sensory stimuli in various ways (FIG. 43-2a). The brain also modifies stimuli to make them more complete, familiar, or logical (FIG. 43-2b).

We can classify sensory receptors based on location of stimuli or on the type of energy they transduce Sensory receptors can be classified according to the location of the stimuli to which they respond. Exteroceptors receive stimuli from the outside environment, enabling an animal to know and explore the world, search for food, find and attract a mate, recognize friends, and detect enemies. Interoceptors are sensory receptors within body organs that detect changes in pH, osmotic pressure, body temperature, and the chemical composition of the blood. You are not usually aware of messages sent to the CNS by these receptors as they work continuously to maintain homeostasis. You become aware of their activity when they signal certain internal conditions such as thirst, hunger, nausea, pain, and orgasm. We can also classify sensory receptors based on the type of energy they transduce (TABLE 43-1). Thermoreceptors respond to heat and cold. Electroreceptors sense differences in electrical potential. Electromagnetic receptors are electroreceptors that are sensitive enough to detect Earth’s magnetic field. Nociceptors (pain receptors) respond to mechanical, temperature, and other stimuli that could be damaging. Mechanoreceptors transduce mechanical energy—touch, pressure, gravity, stretching, and movement. These receptors convert mechanical forces directly into electrical signals. Chemoreceptors transduce the energy of certain chemical compounds, and photoreceptors transduce light energy.

Review ■ ■ ■ ■

What is a receptor potential? What is the function of sensory adaptation? How is sensory information integrated? Name five kinds of sensory receptors based on the type of energy they transduce. What is the specific function of each?

TABLE 43-1

Classification of Receptors by Type of Energy They Transduce

Type of Receptor

Type of Energy Transduced

Examples

Thermoreceptors

Heat

Temperature receptors in blood-sucking insects and ticks; pit organs in pit vipers; nerve endings and receptors in skin and tongues of many animals

Electroreceptors and electromagnetic receptors

Electrical; receptors sense differences in electrical potential; electromagnetic receptors detect magnetic fields

Electrical currents in water used to navigate by many fish and some amphibian species; magnetic fields used for orientation and migration

Nociceptors (pain receptors)

Mechanical; physical force such as strong touch, pressure; heat, temperature extremes; damaging chemicals

Neuron endings in skin and other tissues

Mechanoreceptors

Mechanical; change shape as a result of being pushed or pulled

Tactile receptors (free nerve endings, Merkel cells, Meissner corpuscles, Ruffini endings, Pacinian corpuscles); respond to touch and pressure Proprioceptors; respond to movement and body position Muscle spindles; respond to muscle contraction Golgi tendon organs; respond to stretch of a tendon Joint receptors; respond to movement in ligaments Statocysts in invertebrates; have hair cells that respond to gravity Lateral line organs in fish; detect vibrations in the water; respond to waves and currents Vestibular apparatus Hair cells in saccule and utricle; respond to gravity, linear acceleration Hair cells in semicircular canals; respond to angular acceleration Hair cells in organ of Corti in cochlea; respond to pressure waves (sound)

Chemoreceptors

Specific chemical compounds

Taste buds; olfactory epithelium

Photoreceptors

Light

Eyespots; ommatidia in compound eye of arthropods; rods and cones in retina of vertebrates

43.2 THERMORECEPTORS ■ ■ LEARNING OBJECTIVE 3

Pit organ

Identify three functions of thermoreceptors.

Zig Lesczynski/Animals Animals

Thermoreceptors respond to heat and cold. Mosquitoes, ticks, and other blood-sucking arthropods use thermoreception in their search for an endothermic host. Some have temperature receptors on their antennae that are sensitive to changes of less than 0.5°C. At least two types of snakes—pit vipers and boas—use thermoreceptors to locate their prey (FIG. 43-3). In mammals, which are endothermic, free nerve endings (and perhaps, specialized receptors) in the skin and tongue detect temperature changes in the outside environment. In addition, pain receptors sense extreme temperatures that pose a threat to the body. Thermoreceptors in the hypothalamus detect internal changes in temperature. They also receive and integrate information from receptors on the body surface. The hypothalamus then initiates homeostatic mechanisms that ensure a constant body temperature. FIGURE 43-3 Thermoreception

Review ■

What are the functions of thermoreceptors in mosquitoes, snakes, and mammals?

The pit organ of the bamboo viper (Trimeresurus stejnegeri) is a sense organ located between each eye and nostril. Thermoreceptors in the pit organ can detect the heat emitted by an endothermic animal up to a distance of 1 to 2 m.

43.3 ELECTRORECEPTORS AND ELECTROMAGNETIC RECEPTORS ■ ■ LEARNING OBJECTIVE 4 Describe the functions of electroreceptors and electromagnetic receptors.

Electroreceptors sense differences in electrical potential. Some predatory species of sharks, rays, and bony fishes detect electric fields generated in the water by the muscle activity of their prey. Several groups of fishes have electric organs, specialized muscle or nerve cells that emit electrical signals and receive a feedback signal. In species that produce a weak electric current, electroreceptors may help in navigation. This mechanism is particularly useful in murky water, where visibility and olfaction are poor. Electroreception also appears important in communication, for example, in recognizing a potential mate. Males and females have different frequencies of electric discharge. A few fishes, such as electric eels or electric rays, have electric organs in their heads capable of delivering powerful shocks (up to several hundred volts) that stun prey or predators. Weaker electrical signals are emitted continuously to help in assessing the environment, orienting themselves, and communicating. Some electroreceptors, called electromagnetic receptors, are sensitive enough to detect Earth’s magnetic field. When a shark, ray, or skate swims across one of Earth’s magnetic field lines, its electromagnetic receptors detect changes that occur in electric currents. This information, which is integrated in the brain, helps the animal orient itself. Many other animals, including some insects, bony fishes, amphibians, reptiles, birds, and a few mammals, can also detect magnetic fields and use this information to orient themselves. Migratory birds and sea turtles are well-known examples of animals that use magnetic fields to navigate (discussed further in Chapter 52).

Review ■ ■

What are two functions of electroreceptors? What is the function of electromagnetic receptors?

43.4 NOCICEPTORS ■ ■ LEARNING OBJECTIVE 5

Describe the functions of nociceptors and identify the roles of glutamate and substance P.

Pain receptors, called nociceptors (from the Latin nocere, meaning “to injure”), are free nerve endings (dendrites) of certain sensory neurons found in almost every tissue. Three main types of nociceptors have been identified. Mechanical nociceptors respond to strong tactile stimuli such as cutting, crushing, or pinching. Thermal nociceptors respond to temperature extremes (temperatures

above 45°C or below 5°C). Other nociceptors respond to a variety of damaging stimuli, including certain chemicals. When stimulated, nociceptors transmit signals through sensory neurons to interneurons in the spinal cord. The sensory neurons release the neurotransmitter glutamate, as well as several neuropeptides, including substance P, that enhance and prolong the actions of glutamate. The interneuron transmits the message to the opposite side of the spinal cord and then upward to the thalamus, where pain perception begins. From the thalamus, impulses are sent into the parietal lobes and several other cortical regions, including areas of the limbic system where the emotional aspects of the pain are processed. When pain signals reach the cerebrum, we become fully aware of our pain and can evaluate the situation.

Review ■ ■

What is the function of each main type of nociceptor? What is the function of substance P?

43.5 MECHANORECEPTORS ■ ■ LEARNING OBJECTIVES 6 Compare the functions and mechanisms of action of the following

7 8

mechanoreceptors: tactile receptors, proprioceptors, statocysts, hair cells, and lateral line organs. Compare the structure and function of the saccule and utricle with that of the semicircular canals in maintaining equilibrium. Trace the path taken by sound waves through the structures of the ear and explain how the organ of Corti functions as an auditory receptor.

Mechanoreceptors are activated when they change shape as a result of being mechanically pushed or pulled. They transduce mechanical energy, permitting animals to feel, hear, and maintain balance. These receptors provide information about the shape, texture, weight, and topographic relations of objects in the external environment. Some mechanoreceptors are extremely sensitive. For example, alligators have pressure receptors on their faces that can detect the movement in the surrounding water caused by a single drop of falling water. Certain mechanoreceptors enable an organism to maintain its body position with respect to gravity (for us, head up and feet down; for a dog, dorsal side up and ventral side down; for a tree sloth, ventral side up and dorsal side down). When displaced from its normal position, the animal quickly adjusts its body to reassume its normal orientation. Mechanoreceptors continuously send information to the CNS regarding the position and movements of the body. These receptors also provide information about the operation of internal organs. For example, they inform us about the presence of food in the stomach, feces in the rectum, urine in the bladder, and a fetus in the uterus.

Tactile receptors are located in the skin The simplest mechanoreceptors are free nerve endings in the skin. They detect touch, pressure, vibration, and pain when stimulated

Three types of mechanoreceptors in the skin have encapsulated endings: Meissner corpuscles, Ruffini endings, and Pacinian corpuscles. Meissner corpuscles, located in the upper dermis, are sensitive to light touch and vibration. They adapt quickly to a sustained stimulus. Ruffini endings, which are located in the dermis, inform us of heavy, continuous pressure and stretch of the skin. They adapt very slowly and are thought to provide information about the position of our fingers and hand. Pacinian corpuscles, which are located deep in the dermis, are sensitive to deep pressure that causes rapid movement of the tissues. They are especially sensitive to stimuli that vibrate. The Pacinian corpuscle consists of a neuron ending surrounded by concentric connective tissue layers interspersed with fluid. Compression causes displacement of the layers, which stimulates the axon. Even though the displacement is maintained under steady compression, the receptor potential rapidly falls to zero, and action potentials cease—an excellent example of sensory adaptation.

by objects that contact the body surface. In many invertebrates and vertebrates, tactile (touch) receptors lie at the base of a hair or bristle. These receptors may sense the body’s orientation in space with respect to gravity. They may also detect air and water vibrations and contact with other objects. The tactile receptor is stimulated indirectly when the hair is bent or displaced. A receptor potential develops, and a few action potentials may be generated. This type of receptor responds only when the hair is moving. Even though the hair may be maintained in a displaced position, the receptor is not stimulated unless there is motion. Thousands of specialized tactile receptors are located in mammalian skin (FIG. 43-4). Merkel cells form discs in the deep regions of the epidermis that extend down into the dermis. Merkel cells are important in sensing light touch and enable us to distinguish among the softness of a baby’s skin, the fuzziness of a peach, or the hardness of a book cover. They adapt slowly, permitting us to know that an object continues to touch the skin.

Ruffini endings (pressure)

Free nerve endings (pain)

Meissner corpuscle (touch, vibration)

Hair

Hair receptor (hair movement)

Pacinian corpuscle (deep pressure)

(a) Mechanoreceptors in human skin. This diagrammatic section through the human skin illustrates several types of mechanoreceptors that respond to touch and pressure. Also shown are free nerve endings that respond to pain.

500 μm

Ed Reschke/Peter Arnold, Inc.

Subcutaneous tissue

Dermis

Epidermis

Merkel cells (light touch)

(b) Pacinian corpuscle, a deep pressure receptor. Pressure

Na+

+

Na+

+

Membrane of the Pacinian corpuscle –

Sodium channel closed

– Na+

Sodium channel opens

(c) Pacinian corpuscles sense mechanical forces. Pressure causes Na+ channels to open, depolarizing the axon.

FIGURE 43-4 Animated Sensory receptors in human skin

Proprioceptors help coordinate muscle movement Proprioceptors help animals maintain postural relations—the position of one part of the body with respect to another. Located within muscles, tendons, and joints, proprioceptors continuously respond to tension and movement. With their continuous input, an animal can perceive the positions of its arms, legs, head, and other body parts, along with the orientation of its body as a whole. This information is essential for all forms of locomotion and for all coordinated and skilled movements, from spinning a cocoon to completing a reverse one-and-a-half dive with a twist. With the help of proprioceptors, we carry out activities such as dressing or playing the piano even in the dark. Vertebrates have three main types of proprioceptors: muscle spindles, which detect muscle movement (FIG. 43-5); Golgi tendon organs, which respond to tension in contracting muscles and in the tendons that attach muscle to bone; and joint receptors, which detect movement in ligaments. Impulses from proprioceptors help coordinate the contractions of the several distinct muscles that may be involved in a single movement. Without such receptors, complicated, skilled acts would be impossible. Proprioceptors are also important in maintaining balance. Proprioceptors are probably more numerous and more continually active than any of the other sensory receptors, although

we are less aware of them than of most of the others. As long as the stimulus is present, receptor potentials are maintained (although not at constant magnitude) and action potentials continue to be generated. Proprioceptors continuously send signals to inform the brain about position. The mammalian muscle spindle, one of the more versatile stretch receptors, helps maintain muscle tone. It is a bundle of specialized muscle fibers, with a central region encircled by sensory neuron endings. These neurons continue to transmit signals for a prolonged period, in proportion to the degree of stretch.

Many invertebrates have gravity receptors called statocysts Many invertebrates—from jellyfish to crayfish—have gravity receptors called statocysts (FIG. 43-6). Statocysts are the simplest organs of equilibrium. A statocyst is basically an infolding of the epidermis lined with receptor cells, called sensory hair cells. These hair cells are equipped with sensory hairs (not true hairs). The cavity contains one or more statoliths, tiny granules of loose sand grains or calcium carbonate held together by an adhesive material secreted by cells of the statocyst. Normally, the particles are pulled downward by gravity and stimulate the sensory hair cells.

Golgi tendon organ Muscle fibers

Motor end plates on spindle fibers

Motor end plates Sensory nerve

Nerve ending

Muscle spindle fibers

Sensory hairs Nuclei

Statocyst

Tendon Statolith

Sensory hair cells

FIGURE 43-5 Proprioceptors

FIGURE 43-6 A statocyst

Muscle spindles detect muscle movement. A muscle spindle consists of an elongated bundle of specialized muscle fibers. Golgi tendon organs respond to tension in muscles and their associated tendons.

Many invertebrates have statocysts, receptors that sense gravitational force and provide information about orientation of the body with respect to gravity.

When the animal changes position, the statocyst tilts and the statolith moves in the same direction. This movement bends the sensory hairs. Each sensory hair cell responds maximally when the animal is at a particular position with respect to gravity. The mechanical displacement results in receptor potentials and action potentials that inform the CNS of the change in position. By “knowing” which sensory cells are firing, the animal knows where “down” is and can correct any abnormal orientation. In a classic experiment, the function of the statocyst was demonstrated by substituting iron filings for sand grains in the statocysts of crayfish. The force of gravity was overcome by holding magnets above the animals. The iron filings were attracted upward toward the magnets, and the crayfish began to swim upside down in response to the new information provided by their gravity receptors.

Hair cells are characterized by stereocilia Vertebrate hair cells are mechanoreceptors that detect movement. They are the sensory receptors, located in the lateral line of fishes, that detect water movements (FIG. 43-7). Hair cells also help maintain position and equilibrium and are important in hearing. The surface of a vertebrate hair cell typically has a single long kinocilium and many shorter stereocilia, hairlike projections that increase in length from one side of the hair cell to the other (see Fig. 43-7c). A kinocilium is a true cilium with a 9 + 2 arrangement of microtubules (see Chapter 4). In contrast, stereocilia are not really cilia. They are microvilli that contain actin filaments. Mechanical stimulation of the stereocilia causes voltage changes. Mechanical stimulation in one direction causes depolarization and release of neurotransmitter, whereas mechanical stimulation in the opposite direction results in hyperpolarization. However, hair cells have no axons and do not produce their own action potentials. Rather, they release neurotransmitter that depolarizes associated neurons.

Opening of lateral line canal

Lateral line organs supplement vision in fishes Lateral line organs of fishes and aquatic amphibians detect vibrations in the water. They inform the animal of obstacles in its way and of moving objects such as prey, enemies, and others in its school. Typically, the lateral line organ is a long canal running the length of the body and continuing into the head. The canal is lined with sensory hair cells. The tips of the stereocilia are enclosed by a cupula, a mass of gelatinous material secreted by the hair cells. Pressure waves, currents, and even slight movement in the water cause vibrations in the lateral line organ. As the water moves the cupula, the stereocilia bend. This changes the membrane potential of the hair cell, which may then release neurotransmitter. If an associated sensory neuron is sufficiently stimulated, an action potential may be dispatched to the CNS.

The vestibular apparatus maintains equilibrium When you think of the ear, you probably think of hearing. However, in vertebrates its main function is to help maintain equilibrium. Many vertebrates do not have outer or middle ears, but all have inner ears (FIG. 43-8). The mammalian inner ear includes organs of equilibrium equipped with hair cells that sense the position of the body with respect to gravity. The inner ear is a group of interconnected canals and sacs, called the labyrinth, which includes a membranous labyrinth that fits inside a bony labyrinth. In mammals the membranous labyrinth has two saclike chambers—the saccule and the utricle— and three semicircular canals. Collectively, the saccule, utricle, and semicircular canals are called the vestibular apparatus (see Fig. 43-8b). Destruction of these organs leads to a considerable loss of the sense of equilibrium. A pigeon whose vestibular apparatus has been destroyed cannot fly but in time can relearn how to maintain equilibrium by using visual stimuli.

Cupula

Skin

Stereocilia

Lateral line

Kinocilium

Cupula Hair cell Lateral line nerve

(a) The lateral line organ is a canal that extends the length of the body.

(b) The canal has numerous groupings of hair cells and openings to the outside environment.

FIGURE 43-7 Lateral line organs of fishes These mechanoreceptors detect vibrations in the water, informing the fish of obstacles or moving objects.

Supporting cells

Sensory neuron

(c) The receptor cells are hair cells. Each hair cell has stereocilia and a kinocilium that are covered with a gelatinous cupula. The hair cells respond to waves, currents, and other disturbances in the water.

Auditory bones Malleus

Incus

Stapes

External ear canal

(a) Structure of the human ear, anterior view.

Cochlear nerve

Ampullae Utricle

Semicircular canals Membranous labyrinth Bony labyrinth

Cochlea

Saccule Vestibular nerve

(b) The vestibular apparatus. The saccule, utricle, and semicircular canals together make up the vestibular apparatus. The utricle and saccule are best seen in the posterior view shown here.

FIGURE 43-8 Animated The human ear: hearing and equilibrium

The sensory cells of the saccule and utricle are hair cells similar to those of the lateral line organ (FIG. 43-9). The stereocilia projecting from the hair cells are covered by a gelatinous cupula in which calcium carbonate ear stones, called otoliths, are embedded. The hair cells in the saccule and utricle lie in different planes. Normally, the pull of gravity causes the otoliths to press against the stereocilia, stimulating them to initiate impulses. Sensory neurons at the bases of the hair cells transmit these signals to the brain. When the head is tilted or in linear acceleration (increasing speed

when the body is moving in a straight line), otoliths press on the stereocilia of different cells, deflecting them. Semicircular canals Deflection of the stereocilia toward the kinocilium depolarOval window izes the hair cell. Deflection Vestibular nerve in the opposite direction hyperpolarizes the hair cell. Cochlear Thus, depending on the dinerve rection of movement, the Cochlea hair cells release more or Vestibule less neurotransmitter. The brain interprets the neural Round window messages so the animal is Tympanic membrane aware of its full body position relative to the ground regardless Eustachian tube of how its head is positioned. Information about turning movements, referred to as angular acceleration, is provided by the three semicircular canals. Each canal, a hollow ring connected with the utricle, lies at right angles to the other two and is filled with fluid called endolymph. At one of the openings of each canal into the utricle, there is a small, bulblike enlargement, the ampulla. Within each ampulla lies a clump of hair cells called a crista (pl., cristae), similar to the groups of hair cells in the utricle and saccule. No otoliths are present. The stereocilia of the hair cells of the cristae are stimulated by movements of the endolymph in the canals (FIG. 43-10 on page 922). When the head is turned, there is a lag in the movement of the fluid within the canals. The stereocilia move in relation to the fluid and are stimulated by its flow. This stimulation produces not only the consciousness of rotation but also certain reflex movements in response to it. These reflexes cause the eyes and head to move in a direction opposite that of the original rotation. Because the three canals are in three different planes, head movement in any direction stimulates fluid movement in at least one of the canals. We humans are used to movements in the horizontal plane but not to vertical movements (parallel to the long axis of the upright body). The motion of an elevator or of a ship pitching in a rough sea stimulates the semicircular canals in an unusual way and may cause seasickness or motion sickness, with resultant nausea or vomiting. When a seasick person lies down, the movement stimulates the semicircular canals in a more familiar way and nausea is less likely to occur.

Auditory receptors are located in the cochlea Many arthropods and most vertebrates have sound receptors, but for many of them hearing does not seem to be a sensory priority. It is important in tetrapods, however, and both birds and mammals have a highly developed sense of hearing. Their auditory receptors, located in the cochlea of the inner ear, contain mechanoreceptor hair cells that detect pressure waves.

KEY POINT

The utricle and saccule sense changes in the position of the head relative to gravity, as well as changes in the rate of motion when moving in a straight line.

Stereocilium Kinocilium

Otoliths Cupula

Hair cell Supporting cell

(a) A group of sensory cells in the utricle.

Nerve fiber

Utricle Saccule

(b) Bending the head activates the sensory cells.

FIGURE 43-9 Animated The saccule and utricle: maintaining equilibrium The saccule and utricle sense linear acceleration, allowing us to know our position relative to the ground. Compare the positions of the otoliths and hair cells in (a) with those in (b). Changes in head position cause the force of gravity to distort the cupula, which in turn distorts

The cochlea is a spiral tube that resembles a snail’s shell (FIG. 43-11). If the cochlea were uncoiled, you would see that it consists of three canals separated from one another by thin membranes. The canals come almost to a point at the apex. Two of these canals, the vestibular canal and the tympanic canal, connect at the apex of the cochlea and are filled with a fluid called perilymph. The middle canal, the cochlear duct, is filled with endolymph and contains the auditory organ, the organ of Corti. Each organ of Corti contains about 18,000 hair cells, in rows that extend the length of the coiled cochlea. The stereocilia of each hair cell extend into the cochlear duct. The hair cells rest on the basilar membrane, which separates the cochlear duct from the tympanic canal. Another membrane, the tectorial membrane, overhangs and is in contact with the hair cells.

the stereocilia of the hair cells. The hair cells respond by releasing neurotransmitters. Impulses are transmitted along the vestibulocochlear nerve to the brain.

Sound waves are amplified in the middle ear In terrestrial vertebrates, sound waves in the air are transformed into pressure waves in the cochlear fluid. In the human ear, for example, sound waves pass through the external auditory canal and vibrate the tympanic membrane, or eardrum, which separates the outer ear and the middle ear. The vibrations are transmitted across the middle ear by three tiny bones—the malleus, incus, and stapes (or hammer, anvil, and stirrup, so called because of their shapes). The malleus is in contact with the eardrum, and the stapes is in contact with a thin, membranous region of the cochlea called the oval window. The eustachian tube, which connects the middle ear with the pharynx (throat region), helps equalize pressure between the middle ear and the atmosphere.

The malleus, incus, and stapes act as three interconnected levers that amplify vibrations. A small movement in the malleus causes a larger movement in the incus and a very large movement in the stapes. The vibrations pass through the oval window to the perilymph in the vestibular canal. If sound waves were conducted directly from air to the oval window, much energy would be lost. The middle ear functions to couple sound waves in the air with the pressure waves conducted through the fluid in the cochlea. Because liquids cannot be compressed, the oval window could not cause movement of the fluid in the vestibular canal if there were no escape valve for the pressure. This valve is provided by the membranous round window at the end of the tympanic canal. The pressure wave presses on the membranes separating the three ducts, and is transmitted to the tympanic canal. The pressure wave makes the round window bulge. The movements of the basilar membrane produced by these pulsations cause the stereocilia of the organ of Corti to rub against the overlying tectorial membrane. Stereocilia are bent by their contact with the tectorial membrane. As a result, ion channels in the plasma membrane of the hair cells open. Calcium ions move into the hair cell, causing the release of glutamate. This neurotransmitter binds to receptors on sensory neurons that synapse with each hair cell, leading to depolarization of these sensory neurons. Axons of the sensory neurons join to form the cochlear nerve, a component of the vestibulocochlear nerve (cranial nerve VIII; also referred to as the auditory nerve). We can summarize the sequence of events involved in hearing as follows:

KEY POINT

The semicircular canals detect angular acceleration or deceleration of the head (changes in the position of the head when you are turning it).

Bony portion

Cupula (pushed to left)

Membranous portion

Endolymph flow

Crista Vestibular nerve

Bent stereocilia of hair cells

Direction of body movement

FIGURE 43-10 Semicircular canals and equilibrium When the head changes its rate of rotation, endolymph within the ampulla of the semicircular canal distorts the cupula. The stereocilia of the hair cells bend, increasing the frequency of action potentials in sensory neurons. Information is transmitted to the brain via the vestibular nerve.

sound waves enter external auditory canal ¡ tympanic membrane vibrates ¡ malleus, incus, and stapes amplify vibrations ¡ oval window vibrates ¡ vibrations are conducted through fluid ¡ basilar membrane vibrates ¡ hair cells in organ of Corti are stimulated ¡ cochlear nerve transmits impulses to brain

Sounds differ in pitch, loudness, and tone quality Pitch depends on frequency of sound waves, or number of vibrations per second, and is expressed as hertz (Hz). Low-frequency vibrations result in the sensation of low pitch, whereas highfrequency vibrations result in the sensation of high pitch. Sounds of a given frequency set up resonance waves in the cochlear fluid that cause a particular section of the basilar membrane to vibrate. High frequencies are detected by hair cells located near the base of

the cochlea, whereas low frequencies are sensed by hair cells near the apex of the cochlea. The brain infers the pitch of a sound from the particular hair cells that are stimulated. The human ear typically registers sound frequencies between about 20 and 20,000 Hz (also expressed as 20 kilohertz), and is most sensitive to sounds between 1000 and 4000 Hz (FIG. 43-12 on page 924). Comparing the energy of audible sound waves with the energy of visible light waves, your ear is ten times more sensitive than your eye. Dogs can hear sounds up to 40,000 Hz, and bats can detect frequencies as high as 100,000 Hz. Loud sounds cause resonance waves of greater amplitude (height). The hair cells are more intensely stimulated, and the cochlear nerve transmits a greater number of impulses per second. Variations in tone quality, or timbre, such as when an oboe, a cornet, and a violin play the same note, depend on the number and

KEY POINT

The hair cells in the organ of Corti in the cochlea are the receptors for hearing.

Cochlear nerve, division of the vestibulocochlear (VIII) nerve

Oval window

Organ of Corti Vestibular canal (contains perilymph)

(a) Cross section through the cochlea. The cochlear nerve is shown.

Vestibular membrane Cochlear duct (contains endolymph)

Vestibular canal

Tectorial membrane Organ of Corti

Tectorial membrane

Basilar membrane

Cochlear duct

Stereocilia

Force

Tympanic canal (contains perilymph)

(b) The organ of Corti. Magnified view of the organ of Corti resting on the basilar membrane and covered by the tectorial membrane (cross section).

Hair cell Cochlear nerve Basilar membrane Tympanic canal

Fluid vibrations

(c) How the organ of Corti works. Vibrations transmitted by the malleus, incus, and stapes set the fluid in the tympanic canal in motion. These vibrations (dotted lines) are transmitted to the basilar membrane. As the membrane vibrates, hair cells of the organ of Corti rub against the overlying tectorial membrane. This stimulation depolarizes the hair cells, generating action potentials in the sensory neurons of the cochlear nerve.

FIGURE 43-11 The cochlea, organ of hearing

kinds of overtones, or harmonics, produced. Many hair cells along the basilar membrane are stimulated and vibrate simultaneously, in addition to the main vibration common to all three instruments. Thus, differences in tone quality are recognized in the pattern of the hair cells stimulated.

Deafness may be caused by injury to, or malformation of, either the sound-transmitting mechanism of the outer, middle, or inner ear or the sound-perceiving mechanism of the inner ear. Efferent neurons from the brain stem protect the hair cells of the organ of Corti by dampening their response. Even so, exposure to

3000 Wide, flexible end of basilar membrane

Chemical compounds that are tasted

2000 600 4000

400 1500 Narrow, stiff end of basilar membrane (near oval window)

20

Opening at tip of sensillum

800

200 1000 5000

20,000

7000

Dendrites of the four chemoreceptors

FIGURE 43-12 Distinguishing pitch The basilar membrane is shown here coiled as it would be within the cochlea. Various parts of the basilar membrane vibrate in response to specific frequencies. The narrow, stiff end of the basilar membrane responds to higher frequencies, whereas the wide, flexible end responds to lower frequencies. Pitch depends on frequency of sound waves expressed as hertz (Hz). The human ear is most sensitive to sounds between 1000 and 4000 Hz (darker color).

high-intensity sound, such as heavily amplified music, damages hair cells in the organ of Corti.

Review ■

■ ■ ■

Which sensory receptors permit you to perform actions such as getting dressed or finding your way into bed in the dark? How do hair cells work? What are otoliths, and what is their role in maintaining equilibrium? What is the sequence of events involved in hearing?

43.6 CHEMORECEPTORS

Cuticle Vacuole Hypodermis Supporting cells

Axons

FIGURE 43-13 An insect taste receptor Insects respond to a wide variety of chemical substances found in foods. Each sensillum in the blowfly contains four chemoreceptors. Dendrites that extend to an opening at the tip sense taste. The other end of each chemoreceptor is an axon that transmits sensory signals to the brain.

■ ■ LEARNING OBJECTIVE 9 Compare the structure and function of the receptors of taste and smell, and describe the function of the vomeronasal organ.

Two highly sensitive types of chemoreceptors have evolved for gustation, or taste, and olfaction, or smell. These chemical senses, found throughout the animal kingdom, allow animals to detect chemical substances in food, water, and air. Evaluating such chemical cues allows animals to find food and mates and to avoid predators. Members of the same species use chemoreception to communicate about these critical issues. For terrestrial vertebrates, the sense of smell involves gaseous substances that reach olfactory receptors through the air. The sense of taste depends on materials dissolved in water (or saliva) in the

mouth. For aquatic animals, these distinctions blur. Does a catfish smell or taste the water? The organs of taste and smell in insects are sensory hairs called sensilla (sing., sensillum). Olfactory sensilla are typically found on the antennae. In some insects, including ants, bees, and wasps, taste sensilla are also located on the antennae. Taste sensilla typically project from the mouthparts. In butterflies and flies, they may also project from the end of the leg. In the blowfly, each sensillum contains four chemoreceptors (FIG. 43-13). Dendrites from each cell extend to an opening at the tip, and axons transmit sensory signals to the brain. Recent findings suggest that insects can distinguish among various bitter tastes more precisely than mammals can. This ability is important in dis-

Taste pore

Epithelial cells

Taste bud

Ed Reschke

Papillae

Taste receptor cell

50 μm

(b) LM of a taste bud. Each taste bud is an epithelial capsule containing several taste receptor cells. A single taste receptor cell may respond to more than one category of taste.

(a) Location of taste buds. Taste buds are concentrated in the papillae that cover the surface of the tongue.

Sugar molecule

K+ channel open

Adenylyl cyclase

K+

G protein

Receptor

GTP

ATP

cAMP activates

K+

Protein kinase A 1

2 G protein 3 ATP is Sugar molecule binds with receptor converted to is activated, in plasma membrane cyclic AMP and activates of taste receptor cell. (cAMP). adenylyl cyclase.

K+

K+ channel closes 4

Cyclic AMP activates a protein kinase that closes K+ channels.

(c) Tasting sweet foods. A sugar molecule activates a signal transduction process that leads to an increase in K+ in the cell. The membrane becomes depolarized, resulting in action potentials in sensory neurons.

FIGURE 43-14 Animated Chemoreception: taste

criminating tastes associated with spoiled, bacteria-laden foods or toxins.

Taste receptors detect dissolved food molecules As just described for insects, the ability to discriminate among tastes has survival value. Poisons are generally bitter. In contrast,

foods with a high caloric value often taste sweet. Mammals sense taste with taste buds located in the mouth. In humans, taste buds lie mainly in tiny elevations, or papillae, on the tongue. Each of the 10,000 or so taste buds is an oval epithelial capsule containing about 50 taste receptor cells interspersed with supporting cells (FIG. 43-14). Taste receptor cells are replaced about every ten days. One sensory neuron can innervate several taste buds.

The plasma membrane at the tip of each taste receptor cell has microvilli that extend into a taste pore on the surface of the tongue, where they are bathed in saliva. The taste receptors detect chemical substances dissolved in saliva. Certain molecules, such as those perceived as sweet, activate a signal transduction process involving a G protein (see Fig. 43-14c). According to one model, adenylyl cyclase activity increases, elevating cyclic AMP levels. A protein kinase is activated that phosphorylates and closes K+ channels. This decrease in K+ permeability sets up a depolarizing receptor potential. Action potentials are then generated in sensory neurons that synapse with the taste receptor cell. A second model proposes a pathway involving IP3, which leads to opening of Ca2+ channels (see Chapter 6). When Ca2+ rush into the cell, they also depolarize the taste cell. Traditionally, four basic tastes have been recognized: sweet, sour, salty, and bitter. Recently, a fifth taste, umami, was added. Umami, which is a savory, or meaty taste, is stimulated by glutamate. Umami was first suggested as a basic taste in 1908 by the Japanese physiologist Kikunae Ikeda. In 1908, Ikeda identified glutamate as the compound that triggers the taste. Umami is responsible for the savoriness of certain aged cheeses, soy sauce, anchovy, and some types of seafood. Additional basic tastes have been proposed, including fatty, which is stimulated by fatty acids. Flavor depends on the four or five basic tastes in combination with smell, texture, and temperature. Smell affects flavor because odors pass from the nasal chamber to the mouth. No doubt you have observed that when your nose is congested, food seems to have little “taste.” The taste buds are not affected, but the blockage of nasal passages severely reduces the participation of olfactory reception in the composite sensation of flavor. Awareness of the genetic component of taste can be traced to 1931 when Arthur L. Fox, a chemist at the DuPont Company, synthesized a compound called phenylthiocarbamide (PTC). Some PTC blew into the air and was inhaled by a colleague, who experienced it as very bitter. Because Fox himself could not taste it, he became intrigued and asked other people to taste it. Fox found that about 25% of people were nontasters. Everyone else experienced PTC as bitter. Further research showed that the ability to taste PTC is inherited as a dominant trait. Since that time, taste researchers have found that some tasters are especially sensitive to bitter tastes. In fact, about 25% of the U.S. population are supertasters, 50% are regular tasters, and 25% are nontasters. Supertasters tend to avoid vegetables and fruits that taste bitter, such as broccoli, brussels sprouts, and cabbage. This can be a problem because these foods contain flavonoids and other compounds that may protect against cancer. Continuing investigation of taste preferences and their nutritional consequences may lead to more effective approaches to improving nutrition and to a better understanding of the relative importance of genetics and learning in food selection.

The olfactory epithelium is responsible for the sense of smell Most invertebrates depend on olfaction, the detection of odors, as their main sensory modality. Recent findings indicate that the mechanisms for chemoreception have been highly conserved throughout the animal kingdom.

In terrestrial vertebrates, olfaction occurs in the nasal epithelium. In humans, the olfactory epithelium is found in the roof of the nasal cavity (FIG. 43-15). It contains about 100 million olfactory receptor cells with ciliated tips. The long, nonmotile cilia extend into a layer of mucus on the epithelial surface of the nasal passageway. Receptor molecules on the cilia bind with compounds dissolved in the mucus. The other end of each olfactory receptor cell is an axon that projects directly to the brain. These axons make up the olfactory nerve (cranial nerve I), which extends to the olfactory bulb in the brain. From there, information is transmitted to the olfactory cortex, which is in the limbic system. (Recall that the limbic system is also associated with emotional behavior. Odors are often associated with feelings and memories.) When an odorant, a molecule that can be smelled, binds with a receptor on a cilium of an olfactory receptor cell, a signal transduction process is initiated (see Fig. 43-15c). A G protein is activated, leading to the synthesis of cyclic AMP, which opens nonspecific gated channels in the plasma membrane. These channels permit Na+ and other cations to enter the cell, causing depolarization; this is the receptor potential. The number of odorous molecules determines the intensity, which in turn determines the magnitude of the receptor potential. Genome projects indicate that mammals have about 1000 genes that code for 1000 types of olfactory receptors. However, humans have only about 350 functional olfactory genes. Humans can detect at least seven groups of odors: camphor, musky, floral, peppermint, ether, pungent, and putrid. Each odor consists of several component chemical groups, and each type of receptor may bind with a particular component. The combination of receptors activated determines what odor you perceive. You are capable of perceiving about 10,000 scents. The olfactory receptors respond to remarkably small amounts of a substance. For example, most people can detect ionone, the synthetic substitute for the odor of violets, when it is present in a concentration of only 1 part to more than 30 billion parts of air. Smell is perhaps the sense that adapts most quickly. The olfactory receptors adapt about 50% in the first second or so after stimulation, so even offensively odorous air may seem odorless after only a few minutes.

Many animals communicate with pheromones Animals within many species communicate with one another by releasing pheromones, small, volatile molecules that are secreted into the environment. For example, female moths release pheromones that attract males. Dogs and wolves use pheromones to mark territory. Mammals have specialized chemoreceptor cells that detect pheromones. These cells make up the vomeronasal organ located in the epithelium of the nose (separate from the main olfactory epithelium). The vomeronasal sensory neurons send signals to areas of the hypothalamus, which serves as a link with the endocrine system. (We discuss pheromones further in Chapter 52.)

Review ■

How is signal transduction similar in taste buds and olfactory receptor cells? How is it different?

Sinus

Sinuses

Olfactory bulb

Neurons of the Olfactory tract to brain centers for smell olfactory bulb Olfactory bulb

Olfactory receptor cells

cephalopod mollusks (squids and octopods), arthropods, and vertebrates. Light energy striking a light-sensitive receptor cell containing these pigments triggers chemical changes in the pigment molecules that result in receptor potentials.

Invertebrate photoreceptors include eyespots, simple eyes, and compound eyes

The simplest light-sensitive structures in animals are found in certain cnidarians Nonsensory and flatworms (FIG. 43-16). They are eyeepithelium spots, called ocelli, that detect light but Cilia do not form images. Eyespots are often Wall of bowl-shaped clusters of light-sensitive (b) The olfactory receptor cells. These nasal cavity cells within the epidermis. They may dereceptors are neurons located in the (a) Location of olfactory receptor cells. Smell olfactory epithelium. tect the direction of the light source and depends on thousands of chemoreceptor cells distinguish light intensity. in the roof of the nasal cavity. Effective image formation requires a more complex eye, usually with a lens. A lens is a structure that concentrates light Odorant on a group of photoreceptors. Vision also Adenylyl requires a brain that can interpret the accyclase Na+ channel Na+ tion potentials generated by the photoreG protein closed ceptors. The brain integrates information about movement, brightness, location, position, and shape of the visual stimulus. Two fundamentally different types of eyes Na+ Receptor evolved: the compound eye of arthropods channel ATP cAMP GTP Na+ and the camera eye of vertebrates and cephopens alopod mollusks. The compound eyes of crustaceans and insects differ structurally and func1 Odorant binds to 3 ATP is 4 Gated Na+ 2 G protein is tionally from vertebrate eyes. The surface receptor in plasma channels open, converted to activated and of a compound eye appears faceted, which membrane of leading to cyclic AMP. activates adenylyl means “having many faces,” like a diamond olfactory cell. depolarization. cyclase. (FIG. 43-17). Each facet is the convex cornea (c) The chemistry of smelling. An odorant activates a signal transduction process of one of the eye’s visual units, called om+ permeability. The membrane becomes depolarized, that leads to an increase in Na matidia (sing., ommatidium). The number resulting in transmission of action potentials that transmit information to the brain. of ommatidia varies with the species. For example, each eye of certain crustaceans FIGURE 43-15 Animated Chemoreception: olfaction has only 20 ommatidia, whereas the dragonfly eye has as many as 28,000. The optical part of each ommatidium includes a biconvex lens and a crystalline cone. The lens and crystalline cone focus light 43.7 PHOTORECEPTORS onto photoreceptor cells called retinular cells. These cells have a light-sensitive membrane made up of microvilli that contain rho■ ■ LEARNING OBJECTIVES dopsin. The membranes of several retinular cells may fuse to form 10 Contrast simple eyes, compound eyes, and vertebrate eyes. a central rod-shaped rhabdome that is sensitive to light. 11 Describe the function of each structure of the vertebrate eye. Compound eyes do not perceive form well. Although the lens 12 Compare the two types of photoreceptors, describe the signal transducsystem of each ommatidium is adequate to focus a small inverted tion pathway that is activated when rhodopsin responds to light, and image, there is little evidence that the animal actually perceives summarize the visual pathway. them as images. However, all the ommatidia together produce a Most animals have photoreceptors that use pigments to absorb composite image, or mosaic picture. Each ommatidium, in gatherlight energy. Rhodopsins are the photopigments in the eyes of ing a point of light from a narrow sector of the visual field, is in fact

Terry Ashley/ Tom Stack & Associates

Sensory cell

violet (UV). Accordingly, an insect can see UV light well, and its world of color is very different from ours. Because they reflect UV light to various degrees, flowers that appear identically colored to humans may appear strikingly dissimilar to insects (see Fig. 37-4e).

Vertebrate eyes form sharp images

The position of the eyes in the front of the head of primates and certain birds allows both eyes to focus on the same object. The To cerebral overlap in information they reganglia ceive results in the same visual in500 μm formation striking the two retinas (light-sensitive areas) at the same (a) Planarian flatworms have (b) Structure of the eyespot of time. This binocular vision is imeyespots. a planarian flatworm. portant in judging distance and in depth perception. The vertebrate eye can be FIGURE 43-16 Photoreception: eyespots compared to an old-fashioned The simplest light-sensitive structures are the ocelli, or eyespots, found in certain invertebrates. camera. An adjustable lens can be focused for different distances, and a diaphragm, called the iris, regulates the size of the light sampling a mean intensity from that sector. All these points of light opening, called the pupil (FIG. 43-18). The retina corresponds taken together form a mosaic picture (see Fig. 43-17d). to the light-sensitive film used in a camera. Outside the retina is Arthropod eyes usually adapt to different intensities of light. A the choroid layer, a sheet of cells filled with black pigment that sheath of pigmented cells envelops each ommatidium, and screenabsorbs extra light. This mechanism prevents light from being reing pigments are present in iris cells and in retinular cells. In nocturflected back into the photoreceptors, which would cause blurring nal and crepuscular (active at dusk) insects and many crustaceans, of images. (Cameras are also black on the inside.) The choroid is pigment migrates proximally and distally within each pigmented rich in blood vessels that supply the retina. cell. When the pigment is in the proximal position, each ommaThe outer coat of the eyeball, called the sclera, is a tough, tidium is shielded from its neighbor, and only light entering diopaque, curved sheet of connective tissue that protects the inner rectly along its axis can stimulate the receptors. When the pigment structures and helps maintain the rigidity of the eyeball. On the is in the distal position, light striking at any angle can pass through front surface of the eye, this sheet becomes the thinner, transparseveral ommatidia and stimulate many retinal units. As a result, ent cornea, through which light enters. The cornea serves as a sensitivity is increased in dim light and the eye is protected from fixed lens that focuses light. excessive stimulation in bright light. Pigment migration is under The lens of the eye is a transparent, elastic ball just behind the neural control in insects and under hormonal control in crustairis. It bends the light rays coming in and brings them to a focus ceans. In some species it follows a daily rhythm. on the retina. The lens is aided by the curved surface of the cornea Although the compound eye can form only coarse images, it and by the refractive properties (ability to bend light rays) of the compensates by following flickers to higher frequencies. Flies can liquids inside the eyeball. The anterior cavity between the cornea detect up to about 265 flickers per second. In contrast, the human and the lens is filled with a watery substance, the aqueous fluid. eye can detect only 45 to 53 flickers per second; for us, flickerThe larger posterior cavity between the lens and the retina is filled ing lights fuse above these values, so we see light provided by an with a more viscous fluid, the vitreous body. Both fluids are imordinary bulb as steady and the movement in motion pictures as portant in maintaining the shape of the eyeball by providing an smooth. To an insect, both room lighting and motion pictures internal fluid pressure. must flicker horribly. The insect’s high critical flicker fusion threshAt its anterior margin, the choroid is thick and projects mediold permits immediate detection of even slight movement by prey ally into the eyeball, forming the ciliary body, which consists of or enemy. The compound eye is an important adaptation to the ciliary processes and the ciliary muscle. The ciliary processes are arthropod’s way of life. glandlike folds that project toward the lens and secrete the aqueCompound eyes differ from our eyes in another respect. They ous fluid. are sensitive to wavelengths of light in the range from red to ultraPigment cell

Optic nerve

Lens Ommatidia

Crystalline cone

Crystalline cone stalk

David Scharf/Peter Arnold, Inc.

Iris cell with screening pigment Rhabdome Retinular cell

1 mm

Optic Optic nerve ganglion

(b) Structure of compound eye showing several ommatidia. The eye registers changes in light and shade, permitting the animal to detect movement.

Courtesy of J. L. Gould, Princeton University

(a) SEM of Mediterranean fruit fly (Ceratitis capitata) showing its prominent compound eyes.

Facets

(c) Structure of an ommatidium. The rhabdome is the light-sensitive core of the ommatidium.

pupil. The other is arranged radially and contracts to increase the size of the pupil. Each eye also has six muscles that extend from the surface of the eyeball to various points in the bony socket. These muscles enable the eye as a whole to move and be oriented in a given direction. Cranial nerves innervate the muscles in such a way that the eyes normally move together and focus on the same area. (d) A bee’s-eye view of a flower. This photograph was taken using an optical device to achieve an approximate simulation of how the bee might see this flower. However, the bee would see UV light rather than red, and circles would be vertically elongated ellipses.

FIGURE 43-17 Photoreception: compound eyes

We focus a camera by changing the distance between the lens and the film. The eye has the power of accommodation, the ability to change focus for near or far vision by changing the shape of the lens (FIG. 43-19). This accommodation is accomplished by the ciliary muscle, a part of the ciliary body. To focus on close objects, the ciliary muscle contracts, making the elastic lens rounder. To focus on more distant objects, the ciliary muscle relaxes and the lens assumes a flattened (ovoid) shape. The amount of light entering the eye is regulated by the iris, a ring of smooth muscle that appears as blue, green, gray, or brown depending on the amount and nature of pigment present. The iris consists of two mutually antagonistic sets of muscle fibers. One set is arranged circularly and contracts to decrease the size of the

The retina contains light-sensitive rods and cones The light-sensitive structure in the vertebrate eye is the retina, which lines the posterior two-thirds of the eyeball and covers the choroid. The retina, which is composed of ten layers, contains the photoreceptor cells called, according to shape, rods and cones. In both rod and cone cells, infoldings of the plasma membrane form stacks of membranous discs that contain the photopigments. The discs greatly increase the light-absorbing surface. The human eye has about 125 million rods and 6.5 million cones. Rods function in dim light allowing us to detect shape and movement. They are not sensitive to colors. Because the rods are more numerous in the periphery of the retina, you can see an object better in dim light if you look slightly to one side of it (allowing the image to fall on the rods). Cones respond to light at higher levels of intensity, such as daylight, and they allow us to perceive fine detail. Cones are responsible for color vision; they are differentially sensitive to different wavelengths (colors) of light. In primates and certain hunting birds (such as eagles), the cones are most concentrated in the fovea, a small, depressed area in the center of the retina. The fovea is

Choroid the region of sharpest vision because it has the greatest density of receptor cells and because the retina is thinner there. Retina Sclera Suspensory Light must pass through several layers of connecting neurons ligaments in the retina to reach the rods and cones (FIG. 43-20). This arrangement lets the rods and cones contact a layer of pigmented epithelium that provides retinal, a component of rhodopsin. The retina has five main types of neurons: (1) Photoreceptors (rods and cones) Iris synapse on (2) bipolar cells, which make synaptic contact with Anterior (3) ganglion cells. Two types of lateral interneurons are cavity the (4) horizontal cells, which receive information from the photoreceptor cells and send it to bipolar Lens Pathway cells, and (5) amacrine cells, which receive mesof sages from the bipolar cells and send signals back light Pupil to the bipolar cells or to ganglion cells (FIG. 43-21). Optic Horizontal and amacrine cells help process visual Cornea nerve input within the retina. Interestingly, one group Conjunctiva “Blind of ganglion cells project to (that is, have axons that spot” Fovea Ciliary muscle Ciliary extend to) the suprachiasmatic nucleus, the body’s body Ciliary process main biological clock. These ganglion cells contain Retinal arteries a light-sensitive pigment (melanopsin) that is imporand veins Posterior tant in nonvisual responses to light. cavity Vitreous The axons of the ganglion cells extend across the surbody face of the retina and unite to form the optic nerve. The area where the optic nerve passes out of the eyeball, the optic disk, is FIGURE 43-18 Animated Photoreception: structure known as the blind spot; because it lacks rods and cones, images of the human eye falling on it cannot be perceived. Light passes through the human eye to photoreceptor cells in the retina. In Here is a simplified summary of the visual pathway: this lateral view, the eye is shown partly sectioned along the sagittal plane

to expose its internal structures. light passes through cornea ¡ through aqueous fluid ¡ through lens ¡ through vitreous body ¡ image forms on photoreceptor cells in retina ¡ signal bipolar cells ¡ signal ganglion cells ¡ optic nerve transmits signals to thalamus ¡ integration by visual areas of cerebral cortex

Light activates rhodopsin How is light converted into the neural signals that transmit information about environmental stimuli into pictures in the brain? Rod cells are so sensitive to light that they can respond to a single Ciliary muscles contract; lens “rounds up”

photon. Rhodopsin in the rod cells and some very closely related photopigments in the cone cells are responsible for the ability to see. Rhodopsin consists of opsin, a large protein that is chemically joined with retinal, an aldehyde of vitamin A (see Fig. 3-14). Two isomers of retinal exist: the cis form, which is folded, and the trans form, which is straight. In the dark, opsin binds to retinal in the cis form. Cyclic GMP (cyclic guanosine monophosphate, a molecule similar to cyclic AMP) opens nonspecific channels that permit passage of Na+ and other cations into the rod cell (FIG. 43-22). This process depolar-

Retina

Lens Fovea

Cornea Suspensory ligaments

Optic nerve

(a) Near vision

FIGURE 43-19 Animated Accommodation How the eye changes focus for (a) near and (b) distant vision.

Ciliary muscles relax; lens becomes flatter

(b) Distant vision

Cone

KEY POINT

The rods and cones in the retina are the photoreceptors in vertebrates. Rod

Retina

Discs

Optic nerve

Vitreous body

Ganglion cell Bipolar cell Retina

Choroid layer & sclera

Horizontal cell Bipolar cell

Front of retina

Amacrine cell

Optic nerve fibers

Cone cell

(a) Neurons in the retina. The elaborate interconnections among the various layers of neurons in the retina allow them to interact and influence one another.

Pigmented epithelium

Ganglion cell

Rod cell

Light

To optic nerve

© Lennart Nilsson/from The Incredible Machine, p. 279

FIGURE 43-21 Neural pathway in the retina The photoreceptor cells (rods and cones) are located in the back of the retina. They transmit information to bipolar cells. The bipolar cells synapse with ganglion cells. The horizontal and amacrine cells are important in local processing of visual input within the retina. Axons of ganglion cells make up the optic nerve.

10 μm

(b) SEM of rods and cones. The red, elongated structures are rods. The two shorter, thicker, yellow structures are cones. The rods function in dim light and allow you to see shapes and movement. The cones allow you to view the world in color.

FIGURE 43-20 Animated Organization of the retina

izes the rod cell, which then releases the neurotransmitter glutamate. The glutamate hyperpolarizes the membrane of the bipolar cell so it does not transmit messages. Note that the photoreceptor differs from other neurons in that the ion channels in its membrane are normally open; it is depolarized and continuously releases neurotransmitter. The steady flow of ions into the cell when the environment is dark is called the dark current. Another unusual characteristic of photoreceptors, and of bipolar cells, is that they do not produce action potentials. Their release of neurotransmitter is graded, regulated by the extent of depolarization. When light strikes rhodopsin, light energy is transduced. This process can be considered in three stages:

1. Light activates rhodopsin. Light transforms cis-retinal to transretinal. Rhodopsin changes shape and breaks down into its components, opsin and retinal. This is the light-dependent process in vision. 2. Rhodopsin is part of a signal transduction pathway. When it changes shape, it binds with transducin, a G protein. Transducin activates an esterase that hydrolyzes cyclic GMP (cGMP) to GMP, reducing the concentration of cGMP. 3. The decrease in concentration of cGMP in the rod causes the Na+ channels to close. Fewer cations pass into the rod cell, and it becomes more negative (hyperpolarized). The rod cell then releases less glutamate. Thus, light decreases the number of neural signals from the rod cells.

move from the dark to bright light, your eyes are very sensitive and you may feel “blinded” by the light. Light adaptation occurs as some of the photopigments break down and the sensitivity of your eyes decreases.

Color vision depends on three types of cones Many invertebrates and at least some animals in each vertebrate class have color vision. Color perception depends on cones. Most mammals have two types of cones, but humans and other primates have three types: blue, green, and red. Each contains a slightly different photopigment. Although the retinal portion of the pigment molecule is the same as in rhodopsin, the opsin protein differs slightly in each type of photoreceptor. Each type of cone responds to light within a considerable range of wavelengths but is named for the ability of its pigment to absorb a particular wavelength more strongly than other cones do. For example, red light can be absorbed by all three types of cones, but those cones most sensitive to red act as red receptors. By comparing the relative responses of the three types of cones, the brain can detect colors of intermediate wavelengths. Color blindness occurs when there is a deficiency of one or more of the three types of cones. This is usually an inherited X-linked condition (see Fig. 11-15).

The release of glutamate normally hyperpolarizes the membrane of the bipolar cell, so a decrease in glutamate release results in its depolarization. The depolarized bipolar cell increases its release of a neurotransmitter, which typically stimulates the ganglion cell. When you move from bright daylight to a dark room, you experience “blindness” for a few moments while your eyes adapt. During exposure to bright light, rhodopsin breaks down, decreasing the sensitivity of your eyes. Dark adaptation occurs as enzymes restore rhodopsin to a form in which it can respond to light. In dim light, vision depends on the rods. Conversely, when you

Discs Na+ channel open

Plasma

Na+ membrane

of rod

Photon

Na+

Na+ channel closes

cGMP G protein (transducin)

cGMP

Plasma

Na+ membrane

GMP

of disc

Rod

Rhodopsin (G protein– linked receptor)

Esterase Disc interior

(a) In the dark. Sodium channels are open. The rod cell membrane is depolarized and releases the inhibitory neurotransmitter glutamate (not shown).

(b) In the light. When rhodopsin absorbs light, it changes shape, activating a signal transduction pathway. The concentration of cGMP is reduced. As a result cGMP-gated sodium channels close, hyperpolarizing the membrane. Hyperpolarization results in a decrease in inhibitory signals from the rod cell.

FIGURE 43-22 The biochemistry of vision Inward foldings of the plasma membrane produce a series of discs in each rod cell. In the dark, the rod cell is depolarized; in the light it becomes hyperpolarized.

Integration of visual information begins in the retina The size, intensity, and location of light stimuli determine initial processing in the retina. Rods and cones send signals directly to bipolar cells, and bipolar cells send signals directly to ganglion cells. However, as illustrated in Figure 43-21, photoreceptor cells can also transmit signals to horizontal cells, and bipolar cells can transmit signals to amacrine cells. Ganglion cells are inhibited by amacrine cells and can be inhibited indirectly by horizontal cells as a result of their action on bipolar cells. Thus, the horizontal and amacrine cells integrate signals laterally. Bipolar, horizontal, and amacrine cells combine signals from several photoreceptors. Each ganglion cell has a receptive field, a specific group of photoreceptors that light must strike for the ganglion cell to be stimulated. Ganglion cells are activated or inhibited depending on which photoreceptor cells have been stimulated. The pattern of neuron firing in the retina is very important. The signals sent to ganglion cells depend on spatial patterns and timing of the light striking the retina. Ganglion cells receive signals about specific types of visual stimuli, such as color, brightness, and motion, and transmit this information about the characteristics of a visual image. Ganglion cells produce action potentials, in contrast with rods and cones and most other neurons of the retina, which produce only graded potentials. The retina begins the construction of visual images, but those images are interpreted in the brain. Axons of more than one million ganglion cells form the optic nerves that transmit information to the brain by complex, encoded signals. The optic nerves cross in the floor of the hypothalamus, forming an X-shaped structure, the optic chiasm (FIG. 43-23). Some of the axons of the optic nerves cross over and then extend to the opposite side of the brain. Axons of the optic nerves end in the lateral geniculate nuclei of the thalamus. From there, neurons send signals to the primary visual cortex in the occipital lobe of the cerebrum. The lateral geniculate nucleus controls which information is sent to the visual cortex. Neurons in the reticular activating system are involved in this integration. Information is transmitted from the primary visual cortex to other cortical areas for further processing. Different types of ganglion cells receive information about different aspects of a visual stimulus. As a result, information from each point in the retina is transmitted in parallel by several types of ganglion cells. These cells project to different kinds of neurons in the lateral geniculate nucleus and primary visual cortex. Neurobiologists have not yet discovered all the mechanisms by which the brain makes sense out of the visual information it receives. We know that a large part of the association areas of the

■ ■

Left eye

Right eye

Optic nerve Optic chiasm Optic tract Lateral geniculate nucleus of thalamus

Visual area of occipital lobe

FIGURE 43-23 Animated Neural pathway for transmission of visual information Axons of ganglion cells form the optic nerves. The optic nerves cross, forming the optic chiasm. Many optic nerve fibers end in the lateral geniculate nuclei of the thalamus. From there, signals are sent to the visual area in the occipital lobe. (Brain is viewed from above with overlying structures removed.)

cerebrum are involved in integrating visual input. The neurons of the visual cortex are organized as a map of the external visual field. They are also organized into columns. Each column of neurons receives signals originating from light entering either the right or left eye. Columns are also organized by orientation of light. Neurons in a column receive signals originally triggered by light with the same orientation. However, these signals originate from different locations in the retina.

Review ■ ■

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How does the insect’s compound eye compare with the vertebrate eye? What happens when light strikes rhodopsin? (Include a description of the signal transduction pathway in your answer.) What is the sequence of neural signaling in the retina? What is meant by the statement “Vision happens mainly in the brain”?

S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

43.1 (page 912)

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1 Describe how sensory systems function; include descriptions of sensory receptors, energy transduction, receptor potential, sensory adaptation, and perception.

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Sensory receptors are neuron endings or specialized receptor cells in close contact with neurons. Sense organs consist of sensory receptors and accessory cells. Sensory receptors absorb energy from a stimulus and transduce that energy—energy transduction—into electrical energy. Sensory receptors produce receptor potentials, which are depolarizations

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or hyperpolarizations of the membrane. Receptor potentials are graded responses. Receptor potentials can generate action potentials in sensory neurons, which transmit coded signals to the CNS. Sensory adaptation, the decrease in frequency of action potentials in a sensory neuron even when the stimulus is maintained, decreases response to that stimulus. The brain interprets sensations by converting them to perceptions of stimuli. Sensory perception is the process of selecting, interpreting, and organizing sensory information. Learn more about sensory reception in human skin by clicking on the figure in CengageNOW.

2 Classify sensory receptors according to the location of stimuli to which they respond and according to the types of energy they transduce. Exteroceptors receive stimuli from the outside environment. Interoceptors detect changes within the body, such as changes in pH, osmotic pressure, body temperature, and the chemical composition of the blood. ■ Sensory receptors can be classified according to the types of energy they transduce. Thermoreceptors respond to heat and cold. Electroreceptors sense differences in electrical potential. Some electromagnetic receptors can detect Earth’s magnetic field. Nociceptors (pain receptors) respond to mechanical, temperature, and other stimuli that could be damaging. Mechanoreceptors transduce mechanical energy—touch, pressure, gravity, stretching, and movement. Chemoreceptors transduce certain chemical compounds, and photoreceptors transduce light energy. ■

43.2 (page 915) 3 Identify three functions of thermoreceptors. ■

Thermoreceptors transduce thermal energy; they respond to heat and cold. Some parasites use thermoreceptors to find an endothermic host. Some predators use thermoreceptors to locate endothermic prey. In endothermic animals, thermoreceptors provide cues about body temperature.

43.3 (page 916) 4 Describe the functions of electroreceptors and electromagnetic receptors. Electroreceptors, used by predatory fishes to detect prey, respond to electrical stimuli. ■ Some electroreceptors, called electromagnetic receptors, are sensitive enough to detect Earth’s magnetic field. Some animals use information about magnetic fields to orient themselves. Many migratory animals use magnetic fields to navigate. ■

43.4 (page 916)

Mechanoreceptors transduce mechanical energy, permitting animals to feel, hear, and maintain balance. Mechanoreceptors respond to touch, pressure, gravity, stretch, or movement. They are activated when they change shape as a result of being mechanically pushed or pulled. ■ Tactile receptors in the skin respond to mechanical displacement of hairs or to displacement of the receptor cells themselves. The Pacinian corpuscle is a tactile receptor that responds to touch and pressure. ■ Proprioceptors allow an animal to perceive orientation of its body and the positions of its parts. Muscle spindles, Golgi tendon organs, and joint receptors are proprioceptors that continuously respond to tension and movement. ■ Statocysts, gravity receptors found in many invertebrates, are simple organs of equilibrium. A statocyst consists of a chamber lined with ciliated mechanoreceptors; the chamber contains statoliths, granules of sand grains or calcium carbonate that change position as the animal moves. As statoliths move, sensory hairs bend and different sensory cells are stimulated. This mechanical displacement results in receptor potentials and action potentials that inform the CNS of the change in position. ■ Vertebrate hair cells detect movement; they are found in the lateral line of fishes, the vestibular apparatus, semicircular canals, and cochlea. Each hair cell has a single kinocilium, which is a true cilium, and stereocilia, which are microvilli that contain actin filaments. ■ Lateral line organs supplement vision in fishes and some amphibians by informing the animal of moving objects or objects in its path. 7 Compare the structure and function of the saccule and utricle with that of the semicircular canals in maintaining equilibrium. ■ The vertebrate inner ear consists of a labyrinth of fluid-filled chambers and canals that help maintain equilibrium. The vestibular apparatus in the upper part of the labyrinth consists of the saccule, utricle, and semicircular canals. ■ The saccule and utricle contain otoliths that change position when the head is tilted or when the body is moving in a straight line. The otoliths stimulate hair cells that send signals to the brain, enabling the animal to perceive the direction of gravity. ■ The semicircular canals inform the brain about turning movements. Clumps of hair cells, called cristae, are located within each bulblike enlargement, called an ampulla. Cristae are stimulated by movements of the endolymph, a fluid that fills each canal. 8 Trace the path taken by sound waves through the structures of the ear and explain how the organ of Corti functions as an auditory receptor. ■

Malleus Incus Stapes Semicircular canals

5 Describe the functions of nociceptors and identify the roles of glutamate and substance P. Nociceptors are pain receptors; they are free nerve endings of certain sensory neurons that respond to strong tactile stimuli, temperature extremes, and certain chemicals. Sensory neurons that transmit pain signals release the neurotransmitters glutamate and substance P. Substance P enhances and prolongs the actions of glutamate.

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Cochlea

Tympanic membrane Eustachian tube

43.5 (page 916) 6 Compare the functions and mechanisms of action of the following mechanoreceptors: tactile receptors, proprioceptors, statocysts, hair cells, and lateral line organs.

External ear canal

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■

In terrestrial vertebrates, the organ of Corti within the cochlea contains auditory receptors. Sound waves pass through the external auditory canal and cause the tympanic membrane (eardrum) to vibrate. The ear bones— malleus, incus, and stapes—transmit and amplify the vibrations through the middle ear. Vibrations pass through the oval window to fluid within the vestibular canal. Pressure waves press on the membranes that separate the three ducts of the cochlea. The bulging of the round window serves as an escape valve for the pressure. The pressure waves cause movements of the basilar membrane. These movements stimulate the hair cells of the organ of Corti by rubbing them against the overlying tectorial membrane. Neural impulses are initiated in the dendrites of neurons that lie at the base of each hair cell and are transmitted by the cochlear nerve to the brain.

Each ommatidium has a transparent lens and a crystalline cone that focus light onto receptor cells called retinular cells. ■ Vertebrate eyes form sharp images. 11 Describe the function of each structure of the vertebrate eye. ■ In the human eye, light enters through the cornea, is focused by the lens, and produces an image on the retina. The iris regulates the amount of light that can enter. The internal surface of the eye is covered by the choroid, a black coat that prevents light rays from scattering. ■ The eye has the power of accommodation, the ability to change focus for near or far vision by changing the shape of the lens. The lens is attached to the ciliary muscles. When the ciliary muscles contract, the lens becomes rounder. When they relax, the lens flattens.

Watch the human ear in action by clicking on the figures in CengageNOW.

12 Compare the two types of photoreceptors, describe the signal trans-

43.6 (page 924) 9 Compare the structure and function of the receptors of taste and smell, and describe the function of the vomeronasal organ. Chemoreceptors transduce certain kinds of chemical compounds, allowing gustation, or taste, and olfaction, or smell. ■ In vertebrates, taste receptor cells are specialized epithelial cells in taste buds. The olfactory epithelium contains specialized olfactory cells with axons that extend to the brain as fibers of the olfactory nerves. ■ When a molecule binds with a receptor on a taste receptor cell or an olfactory receptor cell, a signal transduction process involving a G protein is initiated. ■ Receptors that make up the vomeronasal organ, located in the nasal epithelium of mammals, detect pheromones, small, volatile signaling molecules released by animals. ■

Learn more about taste buds and olfactory epithelium by clicking on the figures in CengageNOW.

43.7 (page 927) 10 Contrast simple eyes, compound eyes, and vertebrate eyes. ■

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Photoreceptors transduce light energy and serve as the sensory receptors in eyespots and eyes. Eyespots, or ocelli, found in cnidarians and flatworms, detect light but do not form images. The compound eye of insects and crustaceans consists of visual units called ommatidia, which collectively produce a mosaic image.

Learn more about the human eye and the organization of the retina by clicking on the figures in CengageNOW. duction pathway that is activated when rhodopsin responds to light, and summarize the visual pathway. ■ The retina contains the photoreceptor cells: rods, which function in dim light and form images in black and white; and cones, which function in bright light and permit color vision. ■ The retina also contains bipolar cells that send signals to ganglion cells. Two types of lateral interneurons integrate information: horizontal cells receive signals from the rods and cones and send signals to bipolar cells, and amacrine cells receive signals from bipolar cells and send signals back to bipolar cells and to ganglion cells. ■ When the environment is dark, ion channels in the plasma membranes of rod cells are open and the cells are depolarized; they release glutamate, which hyperpolarizes the membranes of bipolar cells so they do not send signals. ■ When light strikes the photopigment rhodopsin in the rod cells, its retinal portion changes shape and initiates a signal transduction process that involves transducin. This G protein activates an esterase that hydrolyzes cGMP, reducing its concentration. As a result, ion channels close and the membrane becomes hyperpolarized. The rod cells release less glutamate, and fewer signals are transmitted. As a result, bipolar cells become depolarized and release neurotransmitter that stimulates ganglion cells. ■ Axons of the ganglion cells make up the optic nerves. The optic nerves transmit information to the lateral geniculate nuclei in the thalamus. From there, neurons project to the primary visual cortex and then to integration centers in the cerebral cortex. See how vision disorders change the eye by clicking on the figure in CengageNOW.

T E S T YO U R U N D E R S TA N D I N G 1. Which of the following is not true of a sensory receptor? (a) detects a stimulus in the environment (b) converts energy of the stimulus into electrical energy (c) produces a receptor potential (d) produces a graded response (e) interprets sensory stimuli

4. Which of the following are not correctly matched? (a) mechanoreceptors—touch, pressure (b) electroreceptors—voltage (c) photoreceptors—light (d) chemoreceptors—gravity (e) nociceptors—extreme temperature

2. A sensory receptor absorbs energy from some stimulus. The next step is (a) release of neurotransmitter (b) transmission of an action potential (c) energy transduction (d) transmission of a receptor potential (e) sensory adaptation

5. Which of the following is/are associated with informing the brain about turning movements? (a) semicircular canals (b) saccule (c) lymph (d) otoliths (e) utricle

3. Interoceptors (a) help maintain homeostasis (b) are located in the skin (c) are photoreceptors (d) adapt rapidly (e) convert absorbed energy directly into action potentials

6. The auditory receptors are located in the (a) utricle (b) organ of Corti (c) vestibular apparatus (d) saccule (e) semicircular canals 7. Which of the following receptors do not have hair cells? (a) lateral line organ (b) saccule (c) utricle (d) semicircular canals (e) rods

8. In the process of hearing, the basilar membrane vibrates. Which event occurs next? (a) tympanic membrane vibrates (b) bones in middle ear amplify and conduct vibrations (c) cochlear nerve transmits impulses to organ of Corti (d) hair cells in organ of Corti are stimulated (e) vibrations conducted to chemoreceptors 9. In olfaction, (a) pheromones stimulate the taste buds (b) the number of odorous molecules determines the magnitude of the receptor potential (c) chemical signals are converted directly into electrical signals (d) a G protein is activated and leads to the closing of gated channels (e) the concentration of cyclic AMP increases, leading to hyperpolarization of the receptor

10. In the human visual pathway, after light passes through the cornea, it next (a) stimulates ganglion cells (b) passes through the lens (c) sends signals through the optic nerve (d) depolarizes horizontal cells (e) hyperpolarizes rod cells 11. When rhodopsin is activated, it (a) concentrates in bipolar cells (b) transmits more action potentials (c) binds with a G protein (d) sends signals to the lateral geniculate nuclei (e) activates substance P 12. Label the diagram. Refer to Figure 43-18 to check your answers.

Suspensory ligaments

cavity

Pathway of light

Conjunctiva Ciliary body

“Blind spot”

muscle

Retinal arteries and veins

process cavity body

CRITICAL THINKING 1. If all neurons transmit the same type of message, how do you know the difference between sound and light? How are you able to distinguish between an intense pain and a mild one? How are these discriminations adaptive?

4. EVOLUTION LINK. Proteins known as opsins are present in the photoreceptors of all light-sensitive receptors that have been studied. What does this suggest about the relationships of organisms as diverse as amoebas, flatworms, crayfish, and elephants?

2. Connoisseurs can recognize many varieties of cheese or wine by “tasting.” How can this be, when there are only a few types of taste receptors? Organisms have only a few basic tastes. How might that be adaptive?

5. EVOLUTION LINK. Once the ability to produce images evolved, eyes evolved relatively rapidly. Hypothesize why image-forming eyes evolved quickly and are now present in the majority of animal groups.

3. Design a synaptic mechanism by which the same neurotransmitter has an excitatory effect on one bipolar cell in the retina and an inhibitory effect on another.

Additional questions are available in CengageNOW at www.cengage.com/ login.

44

Internal Transport

James D. Mauseth, University of Texas

Colored 3D computed tomography (CT) scan of the human heart and some of its blood vessels. The large cream structures at upper center above the heart are blood vessels that carry blood to and from the heart. The fine blood vessels on the surface of the chambers are coronary arteries, which supply the heart muscle itself with oxygenated blood and nutrients.

KEY CONCEPTS 44.1 Arthropods and most mollusks have an open circu-

44.5 The pacemaker in the sinoatrial node initiates each

latory system in which blood bathes the tissues directly; some invertebrates and all vertebrates have a closed circulatory system in which a heart pumps blood that flows through a continuous circuit of blood vessels.

heartbeat in nonavian reptiles, birds, and mammals; one complete heartbeat makes up a cardiac cycle.

44.6 Blood pressure depends on cardiac output, blood volume, and peripheral resistance.

44.2 Vertebrate blood consists of plasma, which transports nutrients, wastes, gases, and hormones; red blood cells, which are specialized to transport oxygen; white blood cells, which defend the body against disease; and platelets, which function in blood clotting.

44.7 Along with the evolution of lungs, vertebrates devel-

44.3 Three main types of vertebrate blood vessels are ar-

44.8 The vertebrate lymphatic system helps maintain

teries, which carry blood away from the heart; capillaries, which are exchange vessels; and veins, which carry blood back toward the heart.

44.4 During the evolution of terrestrial vertebrates, adaptations in circulatory system structures separated oxygen-rich from oxygen-poor blood. The four-chambered hearts and double circuits of endothermic birds and mammals completely separate oxygen-rich blood from oxygenpoor blood.

oped a double circulation. The pulmonary circulation connects the heart and lungs, whereas the systemic circulation delivers oxygen-rich blood to all of the tissues and organs of the body. fluid homeostasis by returning interstitial fluid to the blood, and defends the body against disease organisms.

44.9 Several modifiable risk factors have been identified for cardiovascular disease, which is the leading cause of death in the world.

M

ost cells require a continuous supply of nutrients and oxygen and removal of waste products. In very small animals, these

metabolic needs are met by simple diffusion, the net movement of particles from a region of higher concentration to a region of lower

44.1 TYPES OF CIRCULATORY SYSTEMS

concentration as a result of random motion. A molecule can diffuse

■ ■ LEARNING OBJECTIVE

1 micrometer (mm) in less than 1 millisecond (msec), so diffusion is

1

adequate over microscopic distances. In invertebrates that are only

Compare and contrast internal transport in animals with no circulatory system, those with an open circulatory system, and those with a closed circulatory system.

a few cells thick, diffusion is an effective mechanism for distributing materials to and from their cells. Consequently, many small, aquatic invertebrates have no circulatory system. Fluid between the cells, called interstitial fluid, or tissue fluid, bathes the cells and provides a medium for diffusion of oxygen, nutrients, and wastes. How much time diffusion requires increases with the square of the distance over which diffusion occurs. A cell that is 10 mm away from its oxygen (or nutrient) supply can receive oxygen by diffusion in about 50 msec, but a cell that is 1000 mm (1 mm) away from its oxygen supply would have to wait several minutes! At this distance, a cell could not survive if it had to depend on diffusion alone. The evolution of specialized circulatory systems allowed animals to increase in size and become many cells thick. A circulatory system reduces the diffusion distance that needed materials must travel. It transports oxygen, nutrients, hormones, and other materials to the interstitial fluid surrounding all the cells and removes metabolic

No specialized circulatory structures are present in sponges, cnidarians (hydras, jellyfish), ctenophores (comb jellies), flatworms, or nematodes (roundworms). In cnidarians, the central gastrovascular cavity serves as a circulatory organ as well as a digestive organ (FIG. 44-1a). As the animal stretches and contracts, body movements stir up the contents of the gastrovascular cavity and help distribute nutrients. The flattened body of the flatworm permits effective gas exchange by diffusion (FIG. 44-1b). Its branched gastrovascular cavity brings nutrients close to all the cells. As in cnidarians, circulation is aided by contractions of the body wall muscles, which move the fluid throughout the gastrovascular cavity. The branching excretory system of planarians provides for internal transport of metabolic wastes that are then expelled from the body. Fluid in the body cavity of nematodes and other pseudocoelomate animals helps circulate materials. Nutrients, oxygen, and wastes dissolve in this fluid and diffuse through it to and from the

wastes. In most animals, a circulatory system interacts with every organ system in the body. The computed tomography (CT) scan shown on the previous page visualizes the human heart and several blood vessels. The human circulatory system, known as the cardiovascular system, is the focus of extensive research because cardiovascular disease is the leading cause of death in the United States

Gastrovascular cavity

and throughout the world. A major risk factor for cardiovascular disease is elevated levels of

Pharynx

cholesterol and low-density lipoprotein (LDL) in the blood. In contrast, high-density lipoprotein

Mouth

(HDL) seems to play a protective role, removing excess cholesterol from the blood and tissues. Cells in the liver and certain other organs bind with the HDL, remove the cholesterol, and use it in synthesizing needed compounds. Researchers have identified a gene in mice that codes for an HDL receptor. When investigators knock out this gene, cholesterol levels more than double. As new receptors

(a) In Hydra and other cnidarians, nutrients circulate through the gastrovascular cavity and come in contact with the inner layer of body cells. Nutrients diffuse the short distance to the outer layer of cells.

(b) In planarian flatworms, flatworms, the branched gastrovascular cavity allows nutrients to come within close proximity of most body cells.

and mechanisms involved in lipid transport and metabolism are discovered, they serve as targets for new drugs and other treatments

FIGURE 44-1 Invertebrates with no circulatory system

for cardiovascular disease.

Red arrows indicate the path of nutrient circulation through the gastrovascular cavity.

cells. Body movements of the animal move the fluid and thus distribute these materials. Larger animals require a circulatory system to efficiently distribute materials. A circulatory system typically has the following components: (1) blood, a connective tissue consisting of cells and cell fragments dispersed in fluid, usually called plasma; (2) a pumping organ, generally a heart; and (3) a system of blood vessels or spaces through which blood circulates. Two main types of circulatory systems are open and closed systems.

Stomach

Ventricle Atrium

Many invertebrates have an open circulatory system Arthropods and most mollusks have an open circulatory system, in which the heart pumps blood into vessels that have open ends. Their blood and interstitial fluid are collectively referred to as hemolymph. This fluid spills out of the open ends of the blood vessels, filling large spaces, called sinuses. The sinuses make up the hemocoel (blood cavity), which is not part of the coelom. (In arthropods and mollusks, the coelom is reduced.) The hemolymph bathes the cells of the body directly. Hemolymph re-enters the circulatory system through openings in the heart (in arthropods) or through open-ended vessels that lead to the gills (in mollusks). In the open circulatory system of most mollusks, the heart has three chambers (FIG. 44-2a). The two atria receive hemolymph from the gills. Then the single ventricle pumps the oxygen-rich hemolymph into blood vessels that conduct it into the large sinuses of the hemocoel. After bathing the body cells, the hemolymph passes into vessels that lead to the gills, where it is recharged with oxygen. The hemolymph then returns to the heart. Some mollusks, as well as arthropods, have a hemolymph pigment, hemocyanin, containing copper that transports oxygen. When oxygenated, hemocyanin is blue and imparts a bluish color to the hemolymph of these animals (the original blue bloods!). In arthropods, a tubular heart pumps hemolymph into arteries, blood vessels that deliver it to the sinuses of the hemocoel (FIG. 44-2b). Hemolymph then circulates through the hemocoel, eventually returning to the pericardial cavity surrounding the heart. Hemolymph enters the heart through ostia, tiny openings equipped with valves that prevent backflow. The rate of hemolymph circulation increases when the animal moves. Thus, when an animal is active and most needs nutrients for fuel, its own movement ensures effective circulation. In crayfish and other crustaceans, gas exchange takes place as hemolymph circulates through the gills. However, an open circulatory system cannot provide enough oxygen to maintain the active lifestyle of insects. Hemolymph mainly distributes nutrients and hormones in insects. Oxygen diffuses directly to the cells through a system of air tubes (tracheae) that make up the respiratory system (see Figure 46-2).

Some invertebrates have a closed circulatory system Annelids, some mollusks (cephalopods), and echinoderms have a closed circulatory system. In them, blood flows through a continuous circuit of blood vessels. The walls of the smallest blood

Gills Hemocoel

(a) In most mollusks, the heart pumps hemolymph into blood vessels that conduct it to the hemocoel. After bathing the cells, the hemolymph enters vessels that conduct it to the gills. The hemolymph is recharged with oxygen and then returned to the heart.

Artery

Ostia

Tubular heart

Hemocoel

(b) In arthropods, a tubular heart pumps hemolymph into arteries that deliver it to the sinuses of the hemocoel. After circulating, hemolymph re-enters the heart through ostia in the heart wall.

FIGURE 44-2 Animated Open circulatory systems In mollusks and arthropods, a heart pumps the blood into arteries that end in sinuses of the hemocoel. Hemolymph circulates through the hemocoel.

vessels, the capillaries, are thin enough to permit diffusion of gases, nutrients, and wastes between blood in the vessels and the interstitial fluid that bathes the cells. Proboscis worms (phylum Nemertea) have a rudimentary closed circulatory system. Their system consists of a complete network of blood vessels but no heart. Blood flow depends on movements of the animal and on contractions in the walls of large blood vessels. Earthworms and other annelids have a closed circulatory system in which two main blood vessels extend lengthwise in the body (FIG. 44-3). The ventral vessel conducts blood posteriorly, and the dorsal vessel conducts blood anteriorly. Dorsal and ventral vessels are connected by lateral vessels in every segment. Branches of the lateral vessels deliver blood to the surface, where it is oxygenated. In the anterior part of the worm, five pairs of contractile blood vessels (sometimes called “hearts”) connect dorsal and ventral vessels. Contractions of these paired vessels and of the dorsal

6. Helps distribute metabolic heat within the body, which helps maintain a constant body temperature in endothermic animals 7. Helps maintain appropriate pH 8. Defends the body against invading microorganisms

Review Dorsal vessel

■

Contractile blood vessels

■

■

How are nutrients and oxygen transported to the body cells in a hydra, flatworm, earthworm, insect, and frog? How does an open circulatory system differ from a closed circulatory system? What are five functions of the vertebrate circulatory system?

44.2 VERTEBRATE BLOOD ■ ■ LEARNING OBJECTIVES Ventral Lateral vessel vessels

FIGURE 44-3 Animated Closed circulatory system of the earthworm Blood circulates through a continuous system of blood vessels. Five pairs of contractile blood vessels deliver blood from the dorsal vessel to the ventral vessel.

vessel, as well as contraction of the body wall muscles, circulate the blood. Earthworms have hemoglobin, the same red pigment that transports oxygen in vertebrate blood. However, their hemoglobin is not contained within red blood cells but is dissolved in the blood plasma. Although other mollusks have an open circulatory system, the fast-moving cephalopods, such as the squid and octopus, require a more efficient means of internal transport. They have a closed system made even more effective by accessory “hearts” at the base of the gills, which speed passage of blood through the gills.

Vertebrates have a closed circulatory system All vertebrates have a ventral, muscular heart that pumps blood into a closed system of blood vessels. Capillaries, the tiniest blood vessels, have very thin walls that permit exchange of materials between blood and interstitial fluid. The vertebrate circulatory system consists of heart, blood vessels, blood, lymph, lymph vessels, and associated organs such as the thymus, spleen, and liver. This system performs several functions: 1. Transports nutrients from the digestive system and from storage depots to each cell 2. Transports oxygen from respiratory structures (gills or lungs) to the cells 3. Transports metabolic wastes from each cell to organs that excrete them 4. Transports hormones from endocrine glands to target tissues 5. Helps maintain fluid balance

2 3

Compare the structure and function of plasma, red blood cells, white blood cells, and platelets. Summarize the sequence of events involved in blood clotting.

In vertebrates, blood consists of a pale yellowish fluid called plasma, in which red blood cells, white blood cells, and platelets are suspended (FIG. 44-4 and TABLE 44-1). In humans the total circulating blood volume is approximately 5 L (5.3 qt) in an adult female and about 5.5 L (5.8 qt) in an adult male. About 55% of the blood volume is plasma. The remaining 45% is made up of blood cells and platelets. Because cells and platelets are heavier than plasma, they can be separated from it by centrifugation. Plasma does not separate from blood cells in the body because the blood is constantly mixed as it circulates in the blood vessels.

Plasma is the fluid component of blood Plasma consists of water (about 92%), proteins (about 7%), salts, and a variety of materials being transported, such as dissolved gases, nutrients, wastes, and hormones. Plasma is in dynamic equilibrium with the interstitial fluid bathing the cells and with the intracellular fluid. As blood passes through the capillaries, substances continuously move into and out of the plasma. Any deviation from its normal composition signals one or more organs to restore homeostasis. Plasma contains several kinds of plasma proteins, each with specific properties and functions: fibrinogen; alpha, beta, and gamma globulins; and albumin. Fibrinogen is one of the proteins involved in the clotting process. When the proteins involved in blood clotting have been removed from the plasma, the remaining liquid is called serum. Alpha globulins include certain hormones and proteins that transport hormones; prothrombin, a protein involved in blood clotting; and high-density lipoproteins (HDL), which transport fats and cholesterol. Beta globulins include other lipoproteins that transport fats and cholesterol, as well as proteins that transport certain vitamins and minerals. The gamma globulin fraction of plasma contains many types of antibodies that provide immunity to diseases such as measles and infectious hepatitis. Purified human gamma globulin is sometimes used to treat certain diseases or to reduce the possibility of contracting a disease.

Whole blood

Plasma

Cell components

Plasma proteins

Lipoproteins

Albumins

Globulins

Fibrinogen

Clotting proteins

Water Salts Dissolved gases Hormones Glucose Wastes

White blood cells (leukocytes)

Granular leukocytes

Agranular leukocytes

7 μm Red blood cells (erythrocytes)

1 to 2 μm Platelets

FIGURE 44-4 Animated Composition of vertebrate blood Blood consists of a fluid plasma in which white blood cells, red blood cells, and platelets are suspended.

Plasma proteins, especially albumins and globulins, help regulate distribution of fluid between plasma and interstitial fluid. Too large to pass readily through the walls of blood vessels, these proteins contribute to the blood’s osmotic pressure, which helps maintain an appropriate blood volume. Plasma proteins (along with the hemoglobin in the red blood cells) are also important acid–base buffers. They help maintain the pH of the blood within a narrow range—near its normal, slightly alkaline pH of 7.4.

10 to 14 μm Neutrophil 15 to 20 μm Monocyte

10 to 14 μm Eosinophil 8 to 10 μm Lymphocyte

Red blood cells transport oxygen Erythrocytes, informally called red blood cells (RBCs), are highly specialized for transporting oxygen. In most vertebrates except mammals, circulating RBCs have nuclei. For example, birds have large, oval, nucleated RBCs. In mammals, the nucleus is ejected from the RBC as the cell develops. Each mammalian RBC is a flexible, biconcave disc 7 to 8 mm in diameter and 1 to 2 mm thick. An internal elastic framework maintains the disc shape and permits the cell to bend and twist as it passes through blood vessels even smaller than its own diameter. Its biconcave shape provides a high ratio of surface area to volume, allowing efficient diffusion of oxygen and carbon dioxide into and out of the cell. In an adult human, about 30 trillion RBCs circulate in the blood, approximately 5 million per mL. Erythrocytes are produced within the red bone marrow of certain bones: vertebrae, ribs, breastbone, skull bones, and long bones. As an RBC develops, it produces large quantities of hemoglobin, the oxygen-transporting pigment that gives vertebrate blood its red color. (Oxygen transport is discussed in Chapter 46.) The lifespan of a human RBC is about 120 days. As blood circulates through the liver and spleen, phagocytic cells remove worn-out RBCs from the

10 to 14 μm Basophil

circulation. These RBCs are then disassembled, and some of their components are recycled. In the human body, more than 2.4 million RBCs are destroyed every second, so an equal number must be produced in the bone marrow to replace them. Red blood cell production is regulated by the hormone erythropoietin, which the kidneys release in response to a decrease in oxygen. Anemia is a deficiency in hemoglobin (often accompanied by a decrease in the number of RBCs). When hemoglobin is insufficient, the amount of oxygen transported is inadequate to supply the body’s needs. An anemic person may complain of feeling weak and may become easily fatigued. Three general causes of anemia are (1) loss of blood from hemorrhage or internal bleeding;

TABLE 44-1

Cell Components of Blood Normal Range

Function

Pathology

Red Blood Cells (RBCs)

Male: 4.2–5.4 million/µL Female: 3.6–5.0 million/µL

Oxygen transport; carbon dioxide transport

Too few: anemia Too many: polycythemia

Platelets

150,000–400,000/µL

Essential for clotting

Clotting malfunctions; bleeding; easy bruising

White Blood Cells (WBCs) Neutrophils

5000–10,000/µL About 60% of WBCs

Phagocytosis

Too many: may be due to bacterial infection, inflammation, myelogenous leukemia Too many: may result from allergic reaction, parasitic infestation

Eosinophils

1%–3% of WBCs

Basophils

1% of WBCs

Lymphocytes

25%–35% of WBCs

Monocytes

6% of WBCs

Play role in allergic responses; release proteins that are toxic to parasitic worms; release compounds important in immune responses May play role in prevention of inappropriate clotting; release histamine, which is important in inflammatory responses Produce antibodies; destroy foreign cells Can differentiate to form macrophages and dendritic cells

(2) decreased production of hemoglobin or red blood cells, as in iron-deficiency anemia or pernicious anemia (which can be caused by vitamin B12 deficiency); and (3) increased rate of RBC destruction—the hemolytic anemias, such as sickle cell anemia (see Chapter 16).

White blood cells defend the body against disease organisms The leukocytes, or white blood cells (WBCs), are specialized to defend the body against harmful bacteria and other microorganisms. Leukocytes are amoeba-like cells capable of independent movement. Some types routinely slip through the walls of blood vessels and enter the tissues. Human blood contains three kinds of granular leukocytes and two types of agranular leukocytes (see Fig. 44-4). Both types are manufactured in the red bone marrow. The granular leukocytes are characterized by large, lobed nuclei and distinctive granules in their cytoplasm. The three varieties of granular leukocytes are the neutrophils, eosinophils, and basophils. Neutrophils, the principal phagocytic cells in the blood, are especially adept at seeking out and ingesting bacteria. They also phagocytose dead cells, a cleanup task that is especially demanding after injury or infection. Most granules in neutrophils contain enzymes that digest ingested material. Eosinophils have large granules that stain bright red with eosin, an acidic dye. Eosinophils increase in number during allergic reactions and during parasitic (for example, tapeworm) infestations. The lysosomes of these WBCs contain enzymes such as peroxidases that degrade the cell membranes of parasitic worms and protozoa. These substances may also injure normal tissue. Eosiniphils also produce substances that promote inflammation. Basophils exhibit deep blue granules when stained with basic dyes. Like eosinophils, these cells play a role in allergic reactions.

Atypical lymphocytes present in infectious mononucleosis; too many may be due to lymphocytic leukemia, certain viral infections May increase in monocytic leukemia and fungal infections

Basophils do not contain lysosomes. Granules in their cytoplasm contain histamine, a substance that dilates blood vessels and makes capillaries more permeable. Basophils release histamine in injured tissues and in allergic responses. Other basophil granules contain heparin, which speeds up fat removal from the blood after you eat a meal high in fat. Heparin is an anticoagulant that may help prevent blood from clotting inappropriately within the blood vessels. Agranular leukocytes lack large, distinctive granules, and their nuclei are rounded or kidney-shaped. Two types of agranular leukocytes are lymphocytes and monocytes. Some lymphocytes are specialized to produce antibodies, whereas others directly attack foreign invaders such as bacteria or viruses (discussed in Chapter 45). Monocytes are the largest WBCs, reaching 20 mm in diameter. During infection, monocytes migrate from the blood into the tissues. They can phagocytose cells and remove toxic molecules. Monocytes can also differentiate into macrophages or dendritic cells. Macrophages are giant scavenger cells that voraciously engulf bacteria, dead cells, and debris. Dendritic cells are also important cells of the immune system. For example, some dendritic cells produce compounds that help the body fight viral infection. We will discuss macrophages and dendritic cells further in Chapter 45. Human blood normally has about 7000 WBCs per mL of blood (only 1 for every 700 RBCs). During bacterial infections the number may rise sharply, so a WBC count is useful in diagnosis. The proportion of each kind of WBC is determined by a differential WBC count. The normal distribution of WBCs is indicated in Table 44-1. Leukemia is a form of cancer in which any one of the various kinds of WBCs multiplies rapidly within the bone marrow. Many of these cells do not mature, and their large numbers crowd out developing RBCs and platelets, leading to anemia and impaired

clotting. A common cause of death from leukemia is internal hemorrhaging, especially in the brain. Another frequent cause of death is infection; although the WBC count may rise dramatically, the cells are immature and abnormal and cannot defend the body against disease organisms. Leukemia is treated with chemotherapy, and sometimes with radiation therapy or bone marrow transplant. Thanks to new treatments, survival rates for individuals with leukemia have increased sharply during the past few years, particularly for children. For example, the survival rate for children younger than five years old with acute lymphocytic leukemia is more than 90%.

Platelets function in blood clotting In most vertebrates other than mammals, the blood contains small, oval, nucleated cells called thrombocytes that function in blood clotting. Mammals have platelets, tiny spherical or disc-shaped bits of cytoplasm that lack nuclei. About 300,000 platelets per mL are present in human blood. Platelets are pinched off from very large cells in the bone marrow. Thus, a platelet is not a whole cell but a fragment of cytoplasm enclosed by a membrane. Platelets function in blood clotting and may also stimulate the immune system. When a blood vessel is cut, it constricts to reduce

1 Injury to

blood vessel

3 Platelets adhere

2 Wall of vessel

4 More permanent

to collagen fibers of damaged vessel wall

contracts

Blood flow decreases

Blood flow

blood loss. Platelets physically patch the break by sticking to the rough, cut edges of the vessel. As platelets begin to gather, they release substances that attract other platelets. The platelets become sticky and adhere to collagen fibers in the blood vessel wall. Within about 5 minutes after injury, they form a platelet plug, or temporary clot. At the same time that the temporary clot forms, a stronger, more permanent clot begins to develop. More than 30 chemical substances interact in this complex process. The series of reactions that leads to clotting is triggered when one of the clotting factors in the blood is activated by contact with the injured tissue. In hemophilia, one clotting factor is absent, as a result of an inherited genetic mutation (see Chapter 16). Prothrombin, a plasma protein manufactured in the liver, requires vitamin K for its production. In the presence of clotting factors, calcium ions, and compounds released from platelets, prothrombin is converted to thrombin. Then thrombin catalyzes the conversion of the soluble plasma protein fibrinogen to an insoluble protein, fibrin. Once formed, fibrin polymerizes, producing long threads that stick to the damaged surface of the blood vessel and form the webbing of the clot. These threads trap blood cells and platelets, which help strengthen the clot. The clotting process is summarized in FIGURE 44-5.

clot forms

Platelet plug

Blood flow decreases

Blood flow ceases

(a) How a blood clot forms when a blood vessel is injured.

Damaged cells and platelets release substances that activate clotting factors

Prothrombin activator

(b) Platelets and a variety of clotting factors are important in blood clotting. The color-enhanced SEM of part of a blood clot shows red blood cells enmeshed in a network of fibrin.

Ca2+

Thrombin

Fibrinogen Ca2+

Fibrin threads (clot)

© Lennart Nilssen/Boehringer Ingelheim International, GmbH

Prothrombin

5 μm

FIGURE 44-5 Blood clotting

KEY POINT

The structure of the wall of each type of blood vessel is adapted for the particular function of the vessel; for example, materials are exchanged through the thin walls of capillaries.

Outer coat (connective tissue)

VEIN

Smooth muscle Lymphatic

Endothelium

Vein Artery ARTERY

Outer coat (connective tissue) Venule

Lymph node

Arteriole Endothelium

Capillary bed

CAPILLARY

Capillaries

Movement of interstitial fluid

(b) Comparison of the walls of an artery, vein, and capillary. All three vessels are lined with endothelium. The capillary wall is only one cell thick, allowing exchange of materials.

Lymph capillaries

Lennart Nilsson, Boehringer Ingelheim International, GmbH

(a) The heart pumps blood into arteries. Blood then flows through arterioles, capillaries, and veins, which return it to the heart. Some plasma leaves the capillaries and becomes interstitial fluid. Lymphatic vessels return excess interstitial fluid to the blood through ducts that lead into large veins in the shoulder region. Blood vessels with oxygen-rich blood are shown in red. Those with oxygen-poor blood are shown in blue.

20 μm

(c) LM of red blood cells passing through capillaries almost in single file.

FIGURE 44-6 Animated Blood and lymphatic vessels

Review ■ ■ ■

What are the functions of the main groups of plasma proteins? What is the function of red blood cells? of neutrophils? What are the major steps in blood clotting?

44.3 VERTEBRATE BLOOD VESSELS ■ ■ LEARNING OBJECTIVE 4 Compare the structure and function of different types of blood vessels, including arteries, arterioles, capillaries, and veins.

The vertebrate circulatory system includes three main types of blood vessels: arteries, capillaries, and veins (FIG. 44-6). An artery carries blood away from a heart chamber toward other tissues. When an artery enters an organ, it divides into many smaller branches called arterioles. The arterioles deliver blood into the microscopic capillaries. After blood circulates through capillary networks within an organ or tissues, capillaries merge to form veins that channel the blood back toward the heart. The wall of an artery or vein has three layers (see Fig. 44-6b). The innermost layer, which lines the blood vessel, consists mainly of endothelium, a tissue that resembles squamous epithelium (see Chapter 39). The middle layer is connective tissue and smooth muscle cells, and the outer coat is connective tissue rich in elastic and collagen fibers. Gases and nutrients cannot pass through the thick walls of arteries and veins. Materials are exchanged between the blood and interstitial fluid bathing the cells through the capillary walls,

Precapillary sphincter

which are only one cell thick. Capillary networks in the body are so extensive that at least one of these tiny vessels is located close to every cell in the body. The total length of all capillaries in the body has been estimated to be more than 96,000 km! Smooth muscle in the arteriole wall can constrict (vasoconstriction) or relax (vasodilation), changing the radius of the arteriole. Such changes help maintain appropriate blood pressure and help control the volume of blood passing to a particular tissue. Changes in blood flow are regulated by the nervous system in response to the metabolic needs of the tissue, as well as by the demands of the body as a whole. For example, when a tissue is metabolizing rapidly, it needs more nutrients and oxygen. During exercise, arterioles within skeletal muscles dilate, increasing by more than tenfold the amount of blood flowing to these muscle tissues. If all your blood vessels dilated at the same time, you would not have enough blood to fill them completely. Normally, your liver, kidneys, and brain receive the lion’s share of blood. However, if an emergency suddenly occurred requiring rapid action, your blood would be rerouted quickly in favor of the heart and muscles. At such a time, the digestive system and kidneys can do with less blood because they are not crucial in responding to the crisis. The small vessels that directly link arterioles with venules (small veins) are metarterioles. The true capillaries branch off from the metarterioles and then rejoin them (FIG. 44-7). True capillaries also interconnect with one another. Wherever a capillary branches from a metarteriole, a smooth muscle cell serves as a precapillary sphincter that opens and closes continuously, directing blood first to one and then to another section of tissue. These precapillary sphincters (along with the smooth muscle in the walls of arteries and arterioles) regulate the blood supply to each organ and its subdivisions.

Precapillary sphincter Metarteriole

Metarteriole

True capillaries Arteriole

True capillaries Venule

(a) Sphincters closed. When a tissue is inactive, only its metarterioles are open.

Arteriole

Venule

(b) Sphincters open. When the tissue becomes active, decreased oxygen tension in the tissue relaxes the precapillary sphincters, and the capillaries open. This process increases the blood supply and thus the delivery of nutrients and oxygen to the active tissue.

FIGURE 44-7 Blood flow through a capillary network As a tissue becomes active, the pattern of blood flow through its capillary networks changes.

Review ■ ■

Compare the functions of arteries, capillaries, and veins. How do arterioles function in maintaining homeostasis?

44.4 EVOLUTION OF THE VERTEBRATE CARDIOVASCULAR SYSTEM ■ ■ LEARNING OBJECTIVE 5

Trace the evolution of the vertebrate cardiovascular system from fish to mammal.

The vertebrate cardiovascular system became modified in the course of evolution, as the site of gas exchange shifted from gills to lungs and as certain vertebrates became active, endothermic animals with higher metabolic rates. The vertebrate heart has one or two atria (sing., atrium), chambers that receive blood returning from the tissues, and one or two ventricles that pump blood into the arteries (FIG. 44-8). Vertebrates in some groups have additional chambers. The fish heart has one atrium and one ventricle. The atrium pumps blood into the ventricle, which pumps blood into a single circuit of blood vessels. Blood is oxygenated as it passes through capillaries in the gills. After blood circulates through the gill capillaries, its pressure is low, so blood passes very slowly to the other organs. The fish’s swimming movements facilitate circulation. Blood returning to the heart has a low oxygen content. A thinwalled sinus venosus receives blood returning from the tissues and pumps it into the atrium. In amphibians, blood flows through a double circuit: the pulmonary circulation and the systemic circulation. Oxygen-rich and oxygen-poor blood are kept somewhat separate. The amphibian heart has two atria and one ventricle. A sinus venosus collects oxygen-poor blood returning from the veins and pumps it into the right atrium. Blood returning from the lungs passes directly into the left atrium. Both atria pump into the single ventricle, but oxygen-poor blood is pumped out of the ventricle before oxygen-rich blood enters it. Blood passes into an artery, the conus arteriosus, equipped with a fold that helps keep the blood separate. Much of the oxygenpoor blood is directed into the pulmonary circulation, which delivers it to the lungs and skin, where it is recharged with oxygen. The systemic circulation delivers oxygen-rich blood into arteries that conduct it to the various tissues of the body. Most nonavian reptiles also have a double circuit of blood flow, made more efficient by a wall that partly divides the ventricles. (In this discussion, birds are considered separately from other reptiles.) Mixing of oxygen-rich and oxygen-poor blood is minimized by the timing of contractions of the left and right sides of the heart and by pressure differences. In crocodilians (crocodiles and alligators), the wall between the ventricles is complete, so the heart consists of two separate atria and two separate ventricles. Thus, a four-chambered heart first evolved among the nonavian reptiles. In contrast to birds and mammals, amphibians and nonavian rep-

tiles do not ventilate their lungs continuously. Therefore, it would be inefficient to pump blood through the pulmonary circulation continuously. The shunts between the two sides of the heart allow blood to be distributed to the lungs as needed. In all birds and mammals (and in crocodilians), the septum (wall) between the ventricles is complete. Biologists hypothesize that the completely divided heart evolved twice during the course of vertebrate evolution; first in the crocodilian–bird clade and then independently in mammals. The septum between the ventricles prevents oxygen-rich blood in the left chamber from mixing with oxygen-poor blood in the right chamber. The conus arteriosus has split and become the base of the aorta (the largest artery) and the pulmonary artery. No sinus venosus is present as a separate chamber, but a vestige remains as the sinoatrial node (the pacemaker). Complete separation of the right and left sides of the heart requires blood to pass through the heart twice each time it tours the body. The complete double circuit allows birds and mammals to maintain relatively high blood pressures in the systemic circulation and modest pressures in the pulmonary circulation. The higher pressure is necessary for efficient circulation of blood through the body. However, the delicate air sacs and capillaries of the lungs would be damaged by this pressure. The double circulatory circuit delivers materials to the tissues rapidly and efficiently. Because their blood contains more oxygen per unit volume and circulates more rapidly than in other vertebrates, the tissues of birds and mammals receive more oxygen. As a result, these animals can maintain a higher metabolic rate and a constant high body temperature even in cold surroundings. The pattern of blood circulation in birds and mammals can be summarized as follows: veins (conduct blood from organs) ¡ right atrium ¡ right ventricle ¡ pulmonary arteries ¡ capillaries in the lungs ¡ pulmonary veins ¡ left atrium ¡ left ventricle ¡ aorta ¡ arteries (conduct blood to organs) ¡ arterioles ¡ capillaries ¡ veins

Review ■

What are some of the major adaptations that occurred in the evolution of the vertebrate cardiovascular system?

44.5 THE HUMAN HEART ■ ■ LEARNING OBJECTIVES 6 Describe the structure and function of the human heart. (Include the 7 8

heart’s conduction system in your answer.) Trace the events of the cardiac cycle and relate normal heart sounds to these events. Define cardiac output, describe how it is regulated, and identify factors that affect it.

Not much bigger than a fist and weighing less than a pound, the human heart is a remarkable organ that beats about 2.5 billion times in an average lifetime, pumping about 300 million L (80 million gallons) of blood. To meet the body’s changing needs,

KEY POINT

The four-chambered heart and double circuit that separates oxygen-rich from oxygen-poor blood are important adaptations that evolved as vertebrates diversified and some became active, terrestrial, endothermic animals.

Pulmonary vein

Atrium

Sinus venosus

Sinus venosus Veins from the body Right atrium

Veins from the body Valve Ventricle Aorta

(a) Fishes. The single atrium and ventricle of the fish heart are part of a single circuit of blood flow.

Semilunar valves

Pulmonary artery Left atrium

Ventricle

Valve

Aorta

Incomplete partition of the ventricle

Conus

(c) Reptiles (except birds). The reptilian heart has two atria and two ventricles. In most nonavian reptiles, the wall separating the ventricles is incomplete, so blood from the right and left chambers mixes to some extent. In crocodiles and alligators, the septum is complete and the heart consists of four separate chambers.

Sinus venosus Veins from the body Left atrium

Pulmonary vein Pulmonary artery Aorta Partition separating atria Atria

Valves

Ventricle

Conus

(b) Amphibians. In amphibians, the heart consists of two atria and one ventricle; blood flows through a double circuit.

AV valve Veins from the body

Pulmonary artery

Right atrium

Aorta

Ventricles Semilunar valves

(d) Birds and mammals. In birds and mammals, the two atria and two ventricles separate oxygen-rich blood from oxygen-poor blood.

FIGURE 44-8 Evolution of the vertebrate cardiovascular system

the heart can vary its output from 5 L to more than 20 L of blood per minute. Your heart is a hollow, muscular organ located in the chest cavity directly under the breastbone. Its wall consists mainly of cardiac muscle attached to a framework of collagen fibers. The pericardium, a tough connective tissue sac, encloses the heart. A smooth layer of endothelium covers the inner surface of the pericardium and the outer surface of the heart. Between these two surfaces is a small pericardial cavity filled with fluid, which reduces friction to a minimum as the heart beats. A wall, or septum, separates the right atrium and ventricle from the left atrium and ventricle. Between the atria, the wall is

known as the interatrial septum; between the ventricles, the interventricular septum. A shallow depression, the fossa ovalis, on the interatrial septum marks the place where an opening, the foramen ovale, was located in the fetal heart. In the fetus, the foramen ovale lets the blood flow directly from right to left atrium, so very little passes to the nonfunctional lungs. A small muscular pouch, called an auricle, lies at the upper surface of each atrium. To keep blood from flowing backward, the heart has valves that close automatically (FIG. 44-9). The valve between the right atrium and right ventricle is called the right atrioventricular (AV) valve, or tricuspid valve. The left AV valve (between the left atrium and left ventricle) is the mitral valve, or bicuspid valve. The AV valves

Superior vena cava

Aorta

Left pulmonary arteries Right pulmonary arteries

Pulmonary artery Pulmonary veins

Pulmonary valve

Left atrium

Right atrium

Mitral valve

Pulmonary veins

Aortic valve

Tricuspid valve

Chordae tendineae (“heartstrings”) Papillary muscles

Right ventricle

Left ventricle

Inferior vena cava

Interventricular septum Aorta

FIGURE 44-9 Animated Section through the human heart showing the valves Note the right and left atria, which receive blood, and the right and left ventricles, which pump blood into the arteries. Arrows indicate the direction of blood flow.

are held in place by stout cords, or “heartstrings,” the chordae tendineae. These cords attach the valves to the papillary muscles that project from the walls of the ventricles. When blood returning from the tissues fills the atria, blood pressure on the AV valves forces them to open into the ventricles, filling the ventricles with blood. As the ventricles contract, blood is forced back against the AV valves, pushing them closed. Contraction of the papillary muscles and tensing of the chordae tendineae prevent the valves from opening backward into the atria. These valves are like swinging doors that open in only one direction. Semilunar valves (named for their flaps, which are shaped like half-moons) guard the exits from the heart. The semilunar valve between the left ventricle and the aorta is the aortic valve, and the one between the right ventricle and the pulmonary artery is the pulmonary valve. When blood passes out of the ventricles, the flaps of the semilunar valves are pushed aside and offer no resistance to blood flow. But when the ventricles are relaxing and filling with blood from the atria, blood pressure in the arteries is higher than in the ventricles. Blood then fills the pouches of the valves, stretching them across the artery so blood cannot flow back into the ventricle.

Each heartbeat is initiated by a pacemaker Horror films sometimes feature a scene in which a heart cut out of a human body continues to beat. Scriptwriters of these tales

actually have some factual basis for their gruesome fantasies because a heart carefully removed from the body can continue to beat for many hours if kept in a nutritive, oxygenated fluid. This is possible because the contractions of cardiac muscle begin within the muscle itself and can occur independently of any nerve supply (FIG. 44-10a). At their ends cardiac muscle cells are joined by dense bands called intercalated discs (FIGS. 44-10b and c). These complex junctions contain gap junctions. Recall from Chapter 5 that in a gap junction, two cells connect through pores. Gap junctions are of great physiological importance because they offer very little resistance to the passage of an action potential. Ions move easily through the gap junctions, allowing the entire atrial (or ventricular) muscle mass to contract as one giant cell. Compared with skeletal muscle that has action potentials typically lasting 1 to 2 milliseconds, the action potentials of cardiac muscle are much longer, several hundred milliseconds. Voltageactivated Ca2+ channels open during depolarization of cardiac muscle fibers. Entry of Ca2+ contributes to the longer depolarization time. Another factor is a type of K+ channel that stays open when the cell is at its resting potential but closes during depolarization, lengthening depolarization time by decreasing the permeability of the membrane to K+. From experiments on isolated cardiac muscle fibers, investigators determined that spontaneous contraction results from the combination of a slow decrease in K+ permeability and a slow increase in Na+ and Ca2+ permeability.

SA node or pacemaker

Left atrium

Right atrium AV node

AV bundle Left ventricle

Purkinje fibers

Right and left branches of AV bundle

Right ventricle

(a) The sinoatrial (SA) node initiates each heartbeat. The action potential spreads through the muscle fibers of the atria, producing atrial contraction. Transmission is briefly delayed at the atrioventricular (AV) node before the action potential spreads through specialized muscle fibers into the ventricles.

SA node ¡ atrial muscle fibers (atria contract) ¡ AV node ¡ AV bundle ¡ right and left bundle branches ¡ Purkinje fibers ¡ conduct impulses to muscle fibers of both ventricles ¡ ventricles contract

Ed Reschke

Nucleus

(b) LM of cardiac muscle.

A specialized conduction system ensures that the heart beats in a regular and effective rhythm. Each beat is initiated by the pacemaker, a small mass of specialized cardiac muscle fibers in the sinoatrial (SA) node. The SA node is located in the posterior wall of the right atrium near the opening of a large vein, the superior vena cava. The action potential in the SA node is triggered mainly by the opening of Ca2+ channels. Ends of the SA node fibers fuse with ordinary atrial muscle fibers, so each action potential spreads through the cardiac muscle fibers of both atria, producing atrial contraction. One group of atrial muscle fibers conducts the action potential directly to the atrioventricular (AV) node, located in the right atrium along the lower part of the septum. Here transmission is delayed briefly so that the atria finish contracting before the ventricles begin to contract. From the AV node, the action potential spreads into specialized muscle fibers that make up the AV bundle. The AV bundle divides, sending branches into each ventricle. Fibers of the bundle branches divide further, eventually forming small Purkinje fibers. These fibers conduct impulses to the muscle fibers of both ventricles.

25 μm

Intercalated discs

As each wave of contraction spreads through the heart, electric currents flow into the tissues surrounding the heart and onto the body surface. By placing electrodes on the body surface on opposite sides of the heart, a physician can amplify and record the electrical activity. The graph produced is called an electrocardiogram (EKG or ECG). Abnormalities in the EKG indicate disorders in the heart or its rhythm. For example, in heart block, impulse transmission is delayed or blocked at some point in the conduction system. Artificial pacemakers can help patients with severe heart block. The pacemaker is implanted beneath the skin and its electrodes connected to the heart. This device provides continuous rhythmic impulses that avoid the block and drive the heartbeat.

The cardiac cycle consists of alternating periods of contraction and relaxation Z-line

Don Fawcett/Visuals Unlimited

Mitochondria

(c) TEM of cardiac muscle.

1 μm

FIGURE 44-10 Animated Conduction system of the heart When action potentials are initiated by the pacemaker cells in the SA node, the gap junctions within the intercalated discs allow the impulses to spread easily from one cardiac muscle fiber to the next within the atria (and within the ventricles). The atria (and then the ventricles) contract as a functional unit.

Each minute the heart beats about 70 times. One complete heartbeat takes about 0.8 second and is referred to as a cardiac cycle. That portion of the cycle in which contraction occurs is known as systole; the period of relaxation is diastole. FIGURE 44-11 shows the sequence of events that occur during one cardiac cycle. When you listen to the heartbeat with a stethoscope, you can hear two main heart sounds, “lub-dup,” which repeat rhythmically. These sounds result from the heart valves closing. When the valves close, they cause turbulence in blood flow that sets up vibrations in the walls of the heart chambers. The first heart sound, “lub,” is lowpitched, not very loud, and fairly long-lasting. It is caused mainly by the closing of the AV (mitral and tricuspid) valves and marks the beginning of ventricular systole. The “lub” sound is quickly followed by the higher-pitched, louder, sharper, and shorter “dup” sound. Heard almost as a quick snap, the “dup” marks the closing of the semilunar valves and the beginning of ventricular diastole.

KEY POINT

The rhythmic changes in electrical activity of the heart are responsible for the cardiac cycle, the pattern of contraction and relaxation that takes place during each heartbeat. Superior vena cava

Aorta Pulmonary artery Semilunar valves

Right atrium

Pulmonary vein

Tricuspid valve

Left atrium

Inferior vena cava

Mitral valve

Period of falling pressure. Blood flows from veins into relaxed atria.

5

1

Atrial systole. Atria contract, pushing blood through open tricuspid and mitral valves into ventricles. Semilunar valves are closed.

Right Left ventricle ventricle

2

Heart sounds

4

3

Beginning of ventricular diastole. Pressure in relaxing ventricles drops below that in arteries. Semilunar valves snap shut, causing second heart sound.

Beginning of ventricular systole. Ventricles contract; pressure within ventricles increases and closes tricuspid and mitral valves, causing first heart sound.

Period of rising pressure. Semilunar valves open when pressure in ventricle exceeds that in arteries. Blood spurts into aorta and pulmonary artery.

FIGURE 44-11 Animated The cardiac cycle The cycle comprises contraction of both atria followed by both ventricles. White arrows indicate the direction of blood flow; dotted lines indicate the change in size as contraction occurs.

The quality of these sounds tells a discerning physician much about the state of the valves. For example, when a valve does not close tightly, blood may flow backward. When the semilunar valves are injured, a soft, hissing noise (“lub-shhh”) known as a heart murmur is heard in place of the normal sound. Valve deformities are sometimes present at birth, or they may result from certain diseases, such as rheumatic fever or syphilis. Diseased or deformed valves can be surgically repaired or replaced with artificial valves. You can measure your heart rate by placing a finger over the radial artery in your wrist or the carotid artery in your neck and counting the pulsations for 1 minute. Arterial pulse is the alternate expansion and recoil of an artery. Each time the left ventricle pumps blood into the aorta, the elastic wall of the aorta expands

to accommodate the blood. This expansion moves in a wave down the aorta and the arteries that branch from it. When this pressure wave passes, the elastic arterial wall snaps back to its normal size.

The nervous system regulates heart rate Although the heart can beat independently, its rate is, in fact, carefully regulated by the nervous and endocrine systems (FIG. 44-12). Sensory receptors in the walls of certain blood vessels and heart chambers are sensitive to changes in blood pressure. When stimulated, the receptors send messages to cardiac centers in the medulla of the brain. These cardiac centers govern two sets of autonomic nerves that pass to the SA node: parasympathetic and sympathetic

to the heart, the heart pumps more blood into the arteries (within physiological limits). Increased venous return stretches the cardiac muscle fibers more and they contract with greater force, increasing stroke volume. Norepinephrine released by sympathetic nerves and epinephrine released by the adrenal glands during stress also increase the force of contraction of cardiac muscle fibers.

Stressors and other stimuli Increases Decreases

Hypothalamus

Cardiac centers in the medulla

Increased Adrenal venous glands return

Sympathetic nerves (accelerator nerves)

Ephinephrine Norepinephrine and Norepinephrine

STROKE VOLUME

X

Cardiac output varies with the body’s need Brain Parasympathetic nerves (vagus)

Increase in body temperature

By multiplying the stroke volume by the number of times the left ventricle beats per minute, we can compute the cardiac output (CO). The CO is the volume of blood pumped by the left ventricle into the aorta in 1 minute. For example, in a resting adult the heart may beat about 72 times per minute and pump about 70 mL of blood with each contraction.

Acetylcholine

HEART RATE

=

CO = stroke volume × heart rate (number of ventricular contractions per minute) = 70 mL/stroke × 72 strokes/min = 5040 mL/min (about 5 L/min)

CARDIAC OUTPUT

FIGURE 44-12 Some factors that influence cardiac output

nerves. Parasympathetic and sympathetic nerves have opposite effects on heart rate (see Fig. 42-17). Parasympathetic nerves release the neurotransmitter acetylcholine, which slows the heart. Acetylcholine slows the rate of depolarization by increasing the membrane’s permeability to K+ (FIG. 44-13a). Sympathetic nerves release norepinephrine, which speeds the heart rate and increases the strength of contraction. Norepinephrine stimulates Ca2+ channel opening during depolarization (FIG. 44-13b). Both norepinephrine and acetylcholine act indirectly on ion channels. They activate a signal transduction process involving a G protein (see Chapter 6). Norepinephrine binds to beta-adrenergic receptors, one of the two main types of adrenergic receptors. These receptors are targeted by beta blockers, drugs that block the actions of norepinephrine on the heart and are used clinically in treating hypertension (high blood pressure) and other types of heart disease. In response to physical and emotional stressors, the adrenal glands release epinephrine and norepinephrine, which speed the heart. An elevated body temperature also increases heart rate. During fever, the heart may beat more than 100 times per minute. As you might expect, heart rate decreases when body temperature is lowered. This is the reason that physicians may deliberately lower a patient’s temperature during heart surgery.

Stroke volume depends on venous return The volume of blood one ventricle pumps during one beat is the stroke volume. Stroke volume depends mainly on venous return, the amount of blood the veins deliver to the heart. According to Starling’s law of the heart, if the veins deliver more blood

Cardiac output varies with changes in either stroke volume or heart rate (see Fig. 44-12). When stroke volume increases, CO increases. The CO varies dramatically with the changing needs of the body. During stress or heavy exercise, the normal heart can increase its CO fivefold so that it pumps 20 to 30 L of blood per minute.

Review ■ ■

■

What sequence of events occurs during cardiac conduction? How is the heart regulated? (Include a description of the actions of acetylcholine and norepinephrine.) What factors influence cardiac output?

44.6 BLOOD PRESSURE ■ ■ LEARNING OBJECTIVE 9 Identify factors that determine and regulate blood pressure, and compare blood pressure in different types of blood vessels.

Blood pressure is the force exerted by the blood against the inner walls of the blood vessels. It is determined by CO, blood volume, and resistance to blood flow (FIG. 44-14a). When CO increases, the increased blood flow causes a rise in blood pressure. When CO decreases, the decreased blood flow causes a fall in blood pressure. If the volume of blood is reduced by hemorrhage or by chronic bleeding, the blood pressure drops. In contrast, an increase in blood volume results in an increase in blood pressure. For example, a high dietary intake of salt causes water retention. This increases blood volume and raises blood pressure. Blood flow is impeded by resistance; when the resistance to flow increases, blood pressure rises. Peripheral resistance is the resistance to blood flow caused by blood viscosity and by friction between blood and the blood vessel wall. In the blood of a healthy person, viscosity remains fairly constant and is only a minor influence on changes in blood pressure. More important is the friction

KEY POINT

Parasympathetic nerves release acetylcholine, which slows the heart; sympathetic nerves release norepinephrine, which increases both the heart rate and the strength of contraction. Parasympathetic neuron 1

Parasympathetic neuron releases acetylcholine.

2

Acetylcholine binds with receptors on plasma membrane of cardiac muscle.

3

Receptor activates G protein.

4

G protein binds with K+ channel, opening it.

5

K+ leaves the cell, hyperpolarizing the membrane. Action potentials occur more slowly.

1

Acetylcholine

5

Acetylcholine receptor 2

K+

K+ channel G protein

Plasma membrane 3

GTP

4

K+

Cardiac muscle

(a) Parasympathetic action on cardiac muscle

1

Sympathetic neuron releases norepinephrine.

2

Norepinephrine binds with receptors on the plasma membrane of cardiac muscle.

Sympathetic neuron

1

Norepinephrine 3

4

5

6

Receptor activates G protein. 2

G protein activates adenylyl cyclase, which converts ATP to cyclic AMP (cAMP).

Cyclic AMP activates protein kinase. Protein kinase phosphorylates Ca2+ channels so that they open more easily when neuron is depolarized. Action potentials occur more rapidly.

β-adrenergic receptor G protein

6

Ca2+ Gate open

Plasma membrane

3

4

GTP

Cardiac muscle

Adenylyl cyclase

Ca2+

cAMP ATP

5

Protein kinase

(b) Sympathetic action on cardiac muscle

FIGURE 44-13 Actions of sympathetic and parasympathetic neurons on cardiac muscle cells Sympathetic and parasympathetic neurons release neurotransmitters that initiate a signal transduction process.

between blood and the blood vessel wall. The length and diameter of a blood vessel determine the amount of surface area in contact with the blood. Blood vessel length does not change, but the diameter, especially of an arteriole, does. A small change in blood ves-

sel diameter causes a big change in blood pressure. Constriction of blood vessels raises blood pressure; dilation lowers blood pressure. Blood pressure in arteries rises during systole and falls during diastole. The National Institutes of Health defines normal blood

100

(a) Blood pressure depends on blood flow and resistance to that flow. A variety of factors affect blood flow and resistance.

Area

Diastolic pressure

60

3000

40

Velocity 2000

20

pressure as a systolic pressure of less than 120 and a diastolic pressure of less than 80. An example of a normal blood pressure, measured in the upper arm with a sphygmomanometer, is 110/73 mm of mercury, abbreviated mm Hg. Systolic pressure is indicated by the first number, diastolic by the second. If you have a blood pressure of 120 to 139 systolic over 80 to 89 diastolic, you are considered prehypertensive, and you need to modify your lifestyle to prevent cardiovascular disease and stroke. Lifestyle changes that will reduce your risk include exercising, losing excess weight, following a heart-healthy diet, reducing salt intake, limiting alcohol, and not smoking. If your systolic pressure consistently measures 140 mm Hg or higher or your diastolic pressure measures 90 mm Hg or higher, you have hypertension, or high blood pressure. Hypertension is a risk factor for atherosclerosis and other cardiovascular disease (discussed later in this chapter). In hypertension, there is usually increased vascular resistance, especially in the arterioles and small arteries. The heart’s workload increases because it must pump against this greater resistance. If this condition persists, the left ventricle enlarges and may deteriorate in function. Heredity, aging, and ethnicity contribute to the development of hypertension.

Arterioles

–10

Aorta

0

Arteries

FIGURE 44-14 Blood pressure Area in graph refers to the surface area of blood vessels in contact with blood.

4000

1000 Vena cavae

Vasoconstriction

5000 80

Veins

Cardiac output

Peripheral resistance

Venules

Blood flow

Blood pressure (mm Hg)

Blood pressure

6000

Systolic pressure

Total cross-sectional area (cm2) of the vascular bed

120

Blood viscosity

Capillaries

Blood volume

0

(b) Blood pressure varies in different types of blood vessels. Systolic and diastolic variations in arterial blood pressures are shown. Note that the venous pressure drops below zero (below atmospheric pressure) near the heart.

smooth muscle. Flow of blood through veins depends on several factors, including skeletal muscle movement, which compresses veins. Most veins larger than 2 mm (0.08 in) in diameter that conduct blood against the force of gravity are equipped with valves to prevent backflow. Such valves usually consist of two cusps formed by inward extensions of the vein wall (FIG. 44-15). When a person stands perfectly still for a long time, as when a soldier stands at attention or a store clerk stands at a cash register, blood tends to pool in the veins. When fully distended, veins can accept no more blood from the capillaries. Pressure in the

Blood pressure varies in different blood vessels As you might guess, blood pressure is greatest in the large arteries, decreasing as blood flows away from the heart and through the smaller arteries and capillaries (FIG. 44-14b). By the time blood enters the veins, its pressure is very low, even approaching zero. Flow rate can be maintained in veins at low pressure because they are low-resistance vessels. Their diameter is larger than that of corresponding arteries, and their walls have little

(a) Resting condition.

(b) Muscles contract. Muscles bulge, compressing veins and forcing blood toward the heart. The lower valve prevents backflow.

FIGURE 44-15 Venous blood flow Contraction of skeletal muscles helps move blood through the veins.

(c) Muscles relax. The vein expands and fills with blood from below. The upper valve prevents backflow.

capillaries increases, and large amounts of plasma are forced out of the circulation through the thin capillary walls. Within just a few minutes, as much as 20% of the blood volume can be lost from the circulation—with drastic effect. Arterial blood pressure falls dramatically, reducing blood flow to the brain. Sometimes the resulting lack of oxygen in the brain causes fainting. This is a protective response because lying in a prone position increases blood supply to the brain. In fact, lifting a person who has fainted to an upright position can result in circulatory shock and even death.

Blood pressure is carefully regulated Each time you get up from a horizontal position, your blood pressure changes. Several mechanisms interact to maintain normal blood pressure so that you do not faint when you get out of bed each morning or change position during the day. When blood pressure decreases, sympathetic nerves to the blood vessels stimulate vasoconstriction, causing the pressure to rise again. Baroreceptors, specialized receptors in the walls of certain arteries and in the heart wall, are sensitive to changes in blood pressure. When an increase in blood pressure stretches the baroreceptors, messages are sent to the cardiac and vasomotor centers in the medulla of the brain. The cardiac center stimulates parasympathetic nerves that slow the heart, lowering blood pressure. The vasomotor center inhibits sympathetic nerves that constrict arterioles; this action causes vasodilation, which also lowers blood pressure. These neural reflexes continuously work in a complementary way to maintain blood pressure within normal limits. Several hormones are also involved in regulating blood pressure (discussed in more detail in Chapters 48 and 49). In response to low blood pressure, the kidneys release renin, which activates the renin–angiotensin–aldosterone pathway. Renin acts on a plasma protein (angiotensinogen), triggering a cascade of reactions that produces the hormone angiotensin II, a powerful vasoconstrictor. Vasoconstriction increases blood pressure, restoring homeostasis. Angiotensin II also acts indirectly to maintain blood pressure by increasing the synthesis and release of the hormone aldosterone by the adrenal glands. Aldosterone increases retention of Na+ by the kidneys, resulting in greater fluid retention and increased blood volume. When the body becomes dehydrated, the osmotic concentration of the blood increases. In response, the posterior lobe of the pituitary gland releases antidiuretic hormone (ADH). ADH increases reabsorption of water in the kidneys (and only a small volume of concentrated urine is produced). Blood volume increases, raising blood pressure and restoring homeostasis. When blood volume increases, the atria of the heart release a hormone called atrial natriuretic peptide (ANP). This hormone increases sodium excretion. As a result, a large volume of dilute urine is produced and blood pressure decreases. Nitric oxide also helps regulate blood pressure by causing vasodilation, thus decreasing blood pressure.

Review ■ ■

What is peripheral resistance? How does it affect blood pressure? Blood pressure is low in capillaries. How does this help retain fluid in the circulation?

44.7 THE PATTERN OF CIRCULATION ■ ■ LEARNING OBJECTIVE 10 Trace a drop of blood through the pulmonary and systemic circulations, naming in sequence each structure through which it passes.

Most vertebrates other than fishes have a double circuit of blood vessels: (1) the pulmonary circulation connects the heart and lungs; and (2) the systemic circulation connects the heart with all the body tissues. You can trace this general pattern of circulation in FIGURE 44-16.

KEY POINT

The right side of the heart receives oxygenpoor blood and pumps it into the pulmonary circulation. The left side of the heart receives oxygen-rich blood from the lungs and pumps it into the systemic circulation.

SYSTEMIC CIRCULATION

Capillary network Brain

Superior vena cava

Carotid artery

PULMONARY CIRCULATION

Pulmonary artery

Left lung

Aorta

LA

RA

Right lung

RV

LV

Pulmonary vein

Pulmonary vein

Inferior vena cava

To lower parts of the body

Capillary network

FIGURE 44-16 Systemic and pulmonary circulation In this highly simplified diagram, red represents oxygen-rich blood and blue represents oxygen-poor blood. Blue screen highlights systemic circulation. Red screen highlights pulmonary circulation.

The pulmonary circulation oxygenates the blood Blood from the tissues returns to the right atrium of the heart. This oxygen-poor blood, loaded with carbon dioxide, is pumped by the right ventricle into the pulmonary circulation. As it emerges from the heart, the large pulmonary trunk branches to form the pulmonary arteries that deliver blood to the lungs. The pulmonary arteries are the only arteries in the body that carry oxygen-poor blood. In the lungs the pulmonary arteries branch into smaller and smaller vessels. These blood vessels give rise to extensive networks of pulmonary capillaries that surround the air sacs of the lungs. As blood circulates through the pulmonary capillaries, carbon dioxide diffuses out of the blood and into the air sacs. Oxygen from the air sacs diffuses into the blood so that by the time it enters the pulmonary veins leading back to the left atrium of the heart, the blood is charged with oxygen. Pulmonary veins are the only veins in the body that carry blood rich in oxygen. In summary, blood flows through the pulmonary circulation in the following sequence: right atrium ¡ right ventricle ¡ pulmonary arteries ¡ pulmonary capillaries (in lungs) ¡ pulmonary veins ¡ left atrium

The systemic circulation delivers blood to the tissues Blood entering the systemic circulation is pumped by the left ventricle into the aorta, the largest artery. Arteries that branch off from the aorta conduct blood to all regions of the body. Some of the principal branches include the coronary arteries to the heart wall itself, the carotid arteries to the brain, the subclavian arteries to the shoulder region, the mesenteric artery to the intestine, the renal arteries to the kidneys, and the iliac arteries to the legs (FIG. 44-17). Each of these arteries gives rise to smaller and smaller vessels, somewhat like branches of a tree that divide until they form tiny twigs. Eventually, blood flows into capillary networks within each tissue or organ. Blood returning from the capillary networks within the brain passes through the jugular veins. Blood from the shoulders and arms drains into the subclavian veins. These veins and others returning blood from the upper portion of the body merge to form a very large vein that empties blood into the right atrium. In humans this vein is called the superior vena cava. The renal veins from the kidneys, iliac veins from the lower limbs, hepatic veins from the liver, and other veins from the lower portion of the body return

Carotid arteries Jugular veins Right subclavian artery Superior vena cava

Axillary artery Right lung Liver Renal vein

Left subclavian vein Aortic arch Left pulmonary vein Left pulmonary artery

Left ventricle Right ventricle

Renal artery Kidney

Inferior vena cava

Abdominal aorta

Common iliac vein

Inferior mesenteric artery

Femoral vein

Common iliac artery External iliac artery Femoral artery

FIGURE 44-17 Blood circulation through some of the principal arteries and veins of the human body Blood vessels carrying oxygen-rich blood are red; blood vessels carrying oxygen-poor blood are blue.

blood to the large inferior vena cava, which delivers blood to the right atrium. As an example of blood circulation through the systemic circuit, let us trace a drop of blood from the heart to the right leg and back to the heart: left atrium ¡ left ventricle ¡ aorta ¡ right common iliac artery ¡ smaller arteries in leg ¡ capillaries in leg ¡ small veins in leg ¡ common iliac vein ¡ inferior vena cava ¡ right atrium

In most fishes, amphibians, and nonavian reptiles, the heart muscle is spongy and receives oxygen directly from the blood as it passes through the heart chambers. However, in birds and mammals the walls of the heart are too thick for nutrients and oxygen to reach all the muscle fibers by diffusion. Instead, the cardiac muscle has its own system of blood vessels. In humans, coronary arteries give rise to a network of capillaries within the heart wall. The coronary veins join to form a large vein, the coronary sinus, which empties directly into the right atrium. Blood almost always travels from artery to capillary to vein to the heart. An exception occurs in the hepatic portal system. Instead of leading directly back to the heart (as most veins do), the hepatic portal vein delivers nutrients from the intestine to the liver. Within the liver, the hepatic portal vein gives rise to an extensive network of tiny blood sinuses. As blood courses through the hepatic sinuses, liver cells remove nutrients and store them. Eventually, liver sinuses merge to form hepatic veins, which deliver blood to the inferior vena cava.

clear, watery fluid formed from interstitial fluid and (2) lymph tissue, a type of connective tissue with large numbers of lymphocytes. Lymph tissue is organized into small masses of tissue called lymph nodes and lymph nodules. The tonsils, thymus gland, and spleen, which consist mainly of lymph tissue, are also part of the lymphatic system. Tiny “dead-end” capillaries of the lymphatic system extend into almost all body tissues (FIG. 44-19). Interstitial fluid enters lymph capillaries, where it becomes lymph. Lymph capillaries join to form larger lymphatics (which you can think of as lymph veins). There are no lymph arteries.

Right lymphatic duct Right subclavian vein Thymus Axillary lymph nodes Lymphatics of breasts

Tonsil Cervical lymph nodes Left subclavian vein Thoracic duct

Spleen

Superficial lymphatics of upper limb

Review ■

■

What sequence of blood vessels and heart chambers would a red blood cell pass through on its way (1) from the inferior vena cava to the aorta and (2) from the renal vein to the renal artery? What is the function of the hepatic portal system? How does its sequence of blood vessels differ from that in most other circulatory routes?

44.8 THE LYMPHATIC SYSTEM

Superficial lymphatics of lower limb

■ ■ LEARNING OBJECTIVE 11 Describe the structure and functions of the lymphatic system.

In addition to the blood circulatory system, vertebrates have an accessory circulatory system, the lymphatic system (FIG. 44-18). The lymphatic system (1) collects and returns interstitial fluid to the blood, (2) launches immune responses that defend the body against disease organisms, and (3) absorbs lipids from the digestive tract. In this section we focus on the first function. We discuss immunity in Chapter 45 and lipid absorption in Chapter 47.

The lymphatic system consists of lymphatic vessels and lymph tissue The lymphatic system consists of (1) an extensive network of lymphatic vessels, or simply lymphatics, that conduct lymph, the

FIGURE 44-18 Animated Human lymphatic system Note that the lymphatic vessels extend into most tissues of the body, but the lymph nodes are clustered in certain regions. The right lymphatic duct drains lymph from the upper right quadrant of the body. The thoracic duct drains lymph from other regions of the body. Both ducts return fluid to the blood.

Arteriole Valve

Venule

Red blood cells

Plasma Connective tissue fibers

Capillary bed Lymph

Plasma

FIGURE 44-19 Animated Lymph capillaries Lymph capillaries drain excess interstitial fluid from the tissues. Note that blood capillaries are connected to vessels at both ends, whereas lymph capillaries (shown in green) are “dead-end streets.” The arrows indicate direction of flow.

Lymph passes from lymph capillaries into lymphatics, which at certain locations empty into lymph nodes. As lymph circulates through the lymph nodes, phagocytes filter out bacteria and other harmful materials. The lymph then flows into lymphatics that conduct it away from the lymph node. Lymphatics from all over the body conduct lymph toward the shoulder region. These vessels join the circulatory system at the base of the subclavian veins by way of ducts: the thoracic duct on the left side and the right lymphatic duct on the right. Tonsils are masses of lymph tissue under the lining of the oral cavity and throat. (When enlarged, the pharyngeal tonsils in back of the nose are called adenoids.) Tonsils help protect the respiratory system from infection by destroying bacteria and other foreign matter that enter the body through the mouth or nose. Unfortunately, tonsils are sometimes overcome by invading bacteria and become the site of frequent infection themselves. Some vertebrates, such as frogs, have lymph “hearts,” which pulsate and squeeze lymph along. However, in mammals the walls of lymphatic vessels themselves pulsate. Valves within the lymphatics prevent lymph from flowing backward. When muscles contract or when arteries pulsate, pressure on the lymphatic vessels increases lymph flow. The rate at which lymph flows is slow and variable. The total lymph flow is about 100 mL per hour—far slower than the 5 L per minute of blood flowing in the cardiovascular system.

stitial fluid, or tissue fluid. It contains no red blood cells or platelets and only a few white blood cells. Its protein content is about onefourth that found in plasma because proteins are too large to pass easily through capillary walls. However, smaller molecules dissolved in the plasma do pass out with the fluid leaving the blood vessels. Thus, interstitial fluid contains glucose, amino acids, other nutrients, and oxygen, as well as a variety of salts. This nourishing fluid bathes all the cells. Three factors are responsible for net filtration pressure, which is the tendency for plasma to leave the blood at the arterial end of a capillary and enter the interstitial fluid (FIG. 44- 20). The main force is hydrostatic pressure, the blood pressure against the capillary wall. This effect is augmented by the osmotic pressure of the interstitial fluid. The principal opposing force is the osmotic pressure of the blood, which is greater than the osmotic pressure of the interstitial fluid and restrains fluid loss from the capillary. At the venous ends of the capillaries, the blood hydrostatic pressure is much lower, and the osmotic pressure of the blood draws fluid back into the capillary. However, not as much fluid is absorbed back into the circulation at the venous ends of capillaries as is filtered out at the arterial ends. Furthermore, protein does not return effectively into the venous capillaries and instead tends to accumulate in the interstitial fluid. These potential problems are so serious that without the lymphatic system, fluid balance in the body would be significantly disturbed within a few hours, and death would occur within about 24 hours. The lymphatic system preserves fluid balance by collecting interstitial fluid, including the protein that accumulates in it, and returning it to the circulation. The walls of the lymph capillaries consist of endothelial cells that overlap slightly. When interstitial fluid accumulates, it presses against these cells, pushing them inward like tiny swinging doors that swing in only one direction. As fluid accumulates within the lymph capillary, these cell doors are pushed closed. Obstruction of lymphatic vessels causes lymphedema, swelling from excessive accumulation of interstitial fluid. Lymphatic vessels can be blocked as a result of injury, inflammation, surgery, or parasitic infection. For example, when a breast is removed (mastectomy) because of cancer, lymph nodes in the armpit are often removed to prevent the spread of cancer cells. The disrupted lymph circulation may cause the patient’s arm to swell. Treatment focuses on minimizing the swelling and controlling discomfort and pain.

Review ■ ■

What are the relationships among plasma, interstitial fluid, and lymph? How does the lymphatic system help maintain fluid balance?

44.9 CARDIOVASCULAR DISEASE ■ ■ LEARNING OBJECTIVE

The lymphatic system plays an important role in fluid homeostasis

12 Trace the progression of atherosclerosis and describe at least five risk

When blood enters a capillary network, it is under rather high pressure, so some plasma is forced out of the capillaries into the tissues. Once it leaves the blood vessels, this fluid is called inter-

Cardiovascular disease is the leading cause of death in the world, accounting for about 30% of deaths each year. In the United States, cardiovascular disease is responsible for about 40% of all deaths,

factors for cardiovascular disease.

KEY POINT 1

More fluid leaves at the arterial end of each capillary than is returned at the venous end. The lymphatic system collects the excess interstitial fluid and returns it to the blood.

At the arterial end of a capillary, blood pressure forces plasma out of the capillary. The osmotic pressure of the blood is an opposing force acting to draw fluid into the blood.

2

Osmotic pressure of the interstitial fluid contributes to the net filtration pressure but does not change much between arterial and venous ends of the capillary.

3

At the venous end of a capillary, fluid enters the blood because blood pressure is much lower.

Venous end of capillary

Arterial end of capillary Hydrostatic pressure (+40)

Osmotic pressure of plasma (−28)

Hydrostatic pressure (+15)

Osmotic pressure of interstitial fluid (+3)

Osmotic pressure of plasma (−28)

Osmotic pressure of interstitial fluid (+3)

(40 + 3) − 28 = +15 Net filtration (pressure drawing fluid out of the capillary)

(15 + 3) − 28 = −10 Net absorption (pressure drawing fluid into the capillary)

FIGURE 44-20 Fluid movement between blood and interstitial fluid The exchange of dissolved materials between blood and interstitial fluid depends on fluid movement between the blood and the interstitial fluid surrounding the capillary. This fluid movement depends on the hydro-

more than all types of cancer combined. According to the Centers for Disease Control and Prevention, heart disease costs the United States more than three billion dollars annually (based on figures for 2010). This cost includes health-care services, medications, and lost productivity. Death often results from some complication of atherosclerosis, thrombosis (clotting), or hypertension (high blood pressure). All three of these conditions are typically involved. Atherosclerosis is a progressive disease in which the walls of certain arteries are damaged, inflamed, and narrowed as a result of lipid deposits in their walls. Although it can affect almost any artery, the disease most often develops in the aorta and in the coronary and cerebral arteries. The earliest events in atherosclerosis begin when excess low-density lipoprotein (LDL), known as “bad” cholesterol, in the blood enters the inner layer of blood vessels. (LDLs will be discussed further in Chapter 47; uptake of LDL by cells was described in Chapter 5.) Recall that the inner layer of a blood vessel is made up of endothelial cells. Excess LDLs in the blood vessel wall become oxidized by free radicals and accumulate between and beneath the endothelial cells. Macrophages phagocytose them and become filled with LDLs. These macrophages are called foam cells. Large

static pressure of the blood and on osmotic pressures. The numbers given are hypothetical and represent millimeters of mercury.

numbers of foam cells lead to the development of fatty streaks, thought to be the initial lesion in the progression of atherosclerosis. Interestingly, fatty streaks have been identified in the arteries of some children and adolescents. Underlying smooth muscle fibers migrate to the inner lining of the artery and multiply there. Connective tissue fibers also proliferate and produce collagen. White blood cells including monocytes and T cells (a type of lymphocyte important in immune responses) move into the region. These cells produce substances that maintain chronic inflammation in the inner layer of the arterial wall. Inflammation, discussed in Chapter 45, is one of the body’s most important responses to infection or injury. The buildup of fatty substances, smooth muscle fibers, macrophages and other cells, collagen, and sometimes, calcium form the atherosclerotic plaque. As the plaque increases in size, remodeling of the artery takes place, so that for a time the size of the lumen (opening) of the blood vessel is maintained. Smooth muscle fibers form a protective fibrous cap, which walls off the plaque from the inner lining of the artery and the blood. Macrophages and other cells eventually die within the plaque, recruiting more macrophages to the area to clean up the debris.

© PHOTOTAKE Inc./ Alamy

Science Photo Library

The progression of atherosclerosis can continue for decades without causing symptoms. However, as atherosclerotic plaque builds, arteries eventually lose their ability to stretch when filled with blood. As the walls thicken, the diameter of the artery decreases, and it gradually becomes occluded (blocked), as shown in FIGURE 44-21. The occluded vessels deliver less blood to the tissues, which then become ischemic, or lacking in blood. Now the tissue is deprived of an adequate oxygen and nutrient supply. In ischemic heart disease, the increased need 500 μm 500 μm for oxygen during exercise or emotional stress results in the pain characteristic of angina pectoris. (b) This artery is severely occluded (a) Normal coronary artery. by the atherosclerotic plaque that Nitroglycerin, in the form of pills or a patch, dilates has built up within its wall. veins, decreasing venous return. Cardiac output is lowered so the heart is not working as hard and FIGURE 44-21 Progression of atherosclerosis requires less oxygen. Nitroglycerin also dilates the LMs of cross sections through two arteries showing changes that take place in atherosclerosis. coronary arteries slightly, allowing more blood to reach the heart muscle. Calcium ion channel blockers types of medications used are statins, medications that lower choare drugs that slow the heart by inhibiting the passage of Ca2+ into cardiac muscle fibers. They also dilate the coronary arteries. lesterol level; aspirin, which reduces the tendency for blood clotMyocardial infarction (MI), or heart attack, is a very serious, ting; and ACE (angiotensin-converting enzyme) inhibitors, drugs often fatal, consequence of cardiovascular disease. MI is the first that decrease blood pressure. By inhibiting ACE, a key enzyme resymptom of atherosclerotic cardiovascular disease for about 65% sponsible for the synthesis of angiotensin II, these drugs decrease of men and almost 50% of women. MI is the leading cause of death blood pressure. (Angiotensin II is a powerful vasoconstrictor and and disability in the United States. MI often results from a sudden also promotes production of the hormone aldosterone, which indecrease in coronary blood supply. If only a small region of cardiac creases sodium reabsorption in the kidneys and thus, retention muscle is affected, the heart may continue to function. Cells in the of water. Angiotensin II and aldosterone are discussed further in region deprived of oxygen die and are replaced by scar tissue. Chapter 48.) The sudden decrease in blood supply associated with MI often Surgical procedures include angioplasty and stent placement, occurs when inflammation causes the fibrous cap over the atheroand coronary bypass surgery. Angioplasty involves inserting a small sclerotic plaque to break open. Platelets adhere to the roughened balloon into an occluded coronary artery. Inflating the balloon arterial wall and initiate clotting. A thrombus is a clot that forms breaks up the plaque in the arterial wall and thus widens the veswithin a blood vessel or within the heart; a thrombus that forms sel. A stent (a mesh tube) left in the artery helps keep the artery within a coronary artery can block a sizable branch of the artery. open. In coronary bypass surgery, veins from another location in Blood flow to a portion of heart muscle may be impeded or comthe patient’s body are grafted around occluded coronary arteries. pletely halted. When a critical region of cardiac muscle is deprived The newly positioned blood vessels allow blood to flow around the of its blood supply, the heart may stop beating. This condition, blocked arteries, restoring adequate blood flow to the affected area. called cardiac arrest, can result in death within a few minutes. Future treatment for damaged hearts may include stem cell theraAn episode of ischemia may trigger a fatal arrhythmia such as pies. Researchers have demonstrated that cardiac stem cells can difventricular fibrillation, a condition in which the ventricles conferentiate into cardiac muscle cells and other types of heart cells. tract very rapidly without actually pumping blood. The pulse may Several major modifiable risk factors for cardiovascular disease stop, and blood pressure may fall precipitously. Ventricular fibrilhave been identified: lation has been linked with about 65% of cardiac arrests. The only 1. Elevated LDL-cholesterol levels in the blood. These levels are aseffective treatment for fibrillation is defibrillation with electric sociated with diets rich in total calories, total fats, saturated shock. The shock appears to depolarize all of the muscle fibers in fats, and cholesterol. the heart so that its timing mechanism can reset. 2. Hypertension. The higher the blood pressure is, the greater the When atherosclerosis develops in the cerebral arteries, less risk. The hormone angiotensin II, which at abnormally high blood is delivered to the brain. This condition can lead to a cerebrolevels is thought to contribute to hypertension, also stimulates vascular accident (CVA), commonly referred to as a stroke. Most inflammation. CVAs are caused by ischemia, typically as a result of thrombosis. 3. Cigarette smoking. The risk of developing atherosclerosis is Current treatment for patients with progressive cardiovascular three to six times as great in smokers as in nonsmokers and is disease includes certain drugs; surgical procedures aimed at restordirectly proportional to the number of cigarettes smoked daily. ing and improving blood flow; and most importantly, education Nicotine constricts blood vessels, and carbon monoxide in the regarding lifestyle changes that reduce risk factors. Among the

smoke damages the inner lining of blood vessels. Components of cigarette smoke cause oxidants to form, which may contribute to the inflammation by increasing oxidation of the altered LDL in the arterial wall. 4. Diabetes mellitus. In this endocrine disorder, glucose is not metabolized normally. The body shifts to fat metabolism, and there is a marked increase in circulating lipids, leading to atherosclerosis. Impaired glucose tolerance is also a risk factor. 5. Physical inactivity. Physical inactivity contributes to more than one-third of the approximately 500,000 deaths related to heart disease in the United States each year. Exercise reduces the concentrations of triacylglycerols (triglycerides) and cholesterol in the blood. (These lipids have been associated with heart disease.) At the same time, exercise increases the con-

■ ■

centration of high-density lipoproteins (HDLs), which protect against heart disease. 6. Overweight and obesity. Being overweight can affect cholesterol levels and increase the risk of hypertension and diabetes. The risk of developing cardiovascular disease also increases with age. Other probable risk factors currently being studied are hereditary predisposition, hormone levels, stress and behavior patterns, and dietary factors.

Review What happens when excess LDL enters the inner layer of arteries? What is ventricular fibrillation? In what way does cigarette smoking contribute to atherosclerosis?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

44.1 (page 938) 1 Compare and contrast internal transport in animals with no circulatory system, those with an open circulatory system, and those with a closed circulatory system. ■ Small, simple invertebrates, such as sponges, cnidarians, and flatworms, depend on diffusion for internal transport. Larger animals require a specialized circulatory system, which typically consists of blood, a heart, and a system of blood vessels or spaces through which blood circulates. In all animals, interstitial fluid, the tissue fluid between cells, brings oxygen and nutrients into contact with cells. ■ Arthropods and most mollusks have an open circulatory system in which blood flows into a hemocoel, bathing the tissues directly. ■ Some invertebrates and all vertebrates have a closed circulatory system in which blood flows through a continuous circuit of blood vessels. ■ The vertebrate circulatory system consists of a muscular heart that pumps blood into a system of blood vessels. The circulatory system transports nutrients, oxygen, wastes, and hormones; helps maintain fluid balance, appropriate pH, and body temperature; and defends the body against disease.

3 Summarize the sequence of events involved in blood clotting. ■

Damaged cells and platelets release substances that activate clotting factors. Prothrombin is converted to thrombin, which catalyzes the conversion of fibrinogen to an insoluble protein, fibrin. Fibrin forms long threads that make up the webbing of the clot.

44.3 (page 945) 4 Compare the structure and function of different types of blood vessels, including arteries, arterioles, capillaries, and veins. ■ Arteries carry blood away from the heart. Arterioles are small arteries that regulate blood pressure by constricting (vasoconstriction) and dilating (vasodilation). Capillaries are the thin-walled exchange vessels through which blood and tissues transfer materials. Veins return blood to the heart. Lymphatic

Vein Artery

Learn more about open and closed circulatory systems by clicking on the figures in CengageNOW.

44.2 (page 940) 2 Compare the structure and function of plasma, red blood cells, white blood cells, and platelets. Plasma consists of water, salts, substances in transport, and plasma proteins, including albumins, globulins, and fibrinogen. ■ Red blood cells (RBCs), also called erythrocytes, transport oxygen and carbon dioxide. Red blood cells produce large quantities of hemoglobin, a red pigment that transports oxygen. ■ White blood cells (WBCs), also called leukocytes, defend the body against disease organisms. Lymphocytes and monocytes are agranular white blood cells; neutrophils, eosinophils, and basophils are granular white blood cells. Monocytes can develop into macrophages and dendritic cells. Platelets patch damaged blood vessels and release substances es■ sential for blood clotting. ■

Learn more about the composition of vertebrate blood by clicking on the figures in CengageNOW.

Lymph node

Capillaries

Lymph capillaries

44.4 (page 946)

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5 Trace the evolution of the vertebrate cardiovascular system from fish to mammal. The vertebrate heart has one or two atria, which receive blood, and one or two ventricles, which pump blood into the arteries. The fish heart consists of a single atrium and ventricle that are part of a single circuit of blood flow. ■ In terrestrial vertebrates, circulatory systems separate oxygenrich from oxygen-poor blood; this allows the higher metabolic rate needed to support an active terrestrial lifestyle. Amphibians have two atria and a single ventricle. Blood flows through a double circuit so that oxygen-rich blood is partly separated from oxygen-poor blood. Most nonavian reptiles have a wall that partly divides the ventricles, minimizing the mixing of oxygen-rich and oxygen-poor blood. ■ The four-chambered hearts of birds and mammals separate oxygenrich blood from oxygen-poor blood. ■

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■

Learn more about determining blood pressure by clicking on the figure in CengageNOW.

44.7 (page 954) 10 Trace a drop of blood through the pulmonary and systemic circulations,

44.5 (page 946) 6 Describe the structure and function of the human heart. (Include the heart’s conduction system in your answer.) The heart is enclosed by a pericardium and has valves that prevent backflow of blood. The valve between the right atrium and ventricle is the right atrioventricular (AV) valve, or tricuspid valve. The valve between the left atrium and ventricle is the mitral valve. Semilunar valves guard the exits from the heart. Cardiac muscle fibers are joined by intercalated discs. ■ The pacemaker in the sinoatrial (SA) node initiates each heartbeat. A specialized electrical conduction system coordinates heartbeats. ■

Learn more about heart anatomy by clicking on the figure in CengageNOW.

naming in sequence each structure through which it passes. The pulmonary circulation connects the heart and the lungs; the systemic circulation connects the heart and the tissues. In the pulmonary circulation, the right ventricle pumps blood into the pulmonary arteries, which go to the lungs. Blood circulates through pulmonary capillaries in the lung and then is conducted to the left atrium by a pulmonary vein. ■ In the systemic circulation, the left ventricle pumps blood into the aorta, which branches into arteries leading to the body organs. After flowing through capillary networks within various organs, blood flows into veins that conduct it to the superior vena cava or inferior vena cava. These large veins return blood to the right atrium. ■ The coronary arteries supply the heart muscle with blood. The hepatic portal system circulates nutrient-rich blood through the liver. ■

See cardiac conduction by clicking on the figure

Learn more about systemic and pulmonary circulation by clicking on the figures in CengageNOW.

in CengageNOW.

7 Trace the events of the cardiac cycle and relate normal heart sounds to these events. ■ One complete heartbeat makes up a cardiac cycle. Contraction occurs during systole. The period of relaxation is diastole. At the beginning of ventricular systole, the closing of the AV valves makes a low-pitched “lub” sound. The closing of the semilunar valves, the beginning of ventricular diastole, makes a short, loud “dup” sound. Watch the cardiac cycle by clicking on the figure in CengageNOW.

8 Define cardiac output, describe how it is regulated, and identify factors that affect it. Cardiac output (CO) equals stroke volume times heart rate. Stroke volume depends on venous return and on neural messages and hormones, especially epinephrine and norepinephrine. According to Starling’s law of the heart, the more blood delivered to the heart by the veins, the more blood the heart pumps. Heart rate is regulated mainly by the nervous system and is influ■ enced by hormones and body temperature. ■

44.6 (page 951) 9 Identify factors that determine and regulate blood pressure, and compare blood pressure in different types of blood vessels. Blood pressure is the force blood exerts against the inner walls of the blood vessel. Blood pressure is greatest in the arteries and decreases as blood flows through the capillaries.

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Blood pressure depends on cardiac output, blood volume, and resistance to blood flow. Peripheral resistance is the resistance to blood flow caused by blood viscosity and by friction between blood and the blood vessel wall. Baroreceptors sensitive to blood pressure changes send messages to the cardiac and vasomotor centers in the medulla of the brain. When informed of an increase in blood pressure, the cardiac center stimulates parasympathetic nerves that slow the heart rate. The vasomotor center inhibits sympathetic nerves that constrict blood vessels. These actions reduce blood pressure. Angiotensin II is a hormone that raises blood pressure. Aldosterone helps regulate salt excretion, which affects blood volume and blood pressure.

44.8 (page 956) 11 Describe the structure and functions of the lymphatic system. ■

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The lymphatic system helps maintain homeostasis of fluids by collecting interstitial fluid and returning it to the blood. The lymphatic system also defends the body against disease and absorbs lipids from the digestive tract. Lymphatic vessels conduct lymph, a clear fluid formed from interstitial fluid, to the thoracic duct and right lymphatic duct in the shoulder region. These ducts return lymph to the blood. Lymph nodes are small masses of tissue that filter bacteria and harmful materials out of lymph. Learn more about the human lymphatic system by clicking on the figure in CengageNOW.

44.9 (page 957) 12 Trace the progression of atherosclerosis and describe at least five risk factors for cardiovascular disease. Atherosclerosis progresses as the inner walls of certain arteries ■ become inflamed and blocked by the accumulation of fatty deposits and cells that mediate inflammation. The blood supply of tissues served by these vessels becomes increasingly compromised. Risk factors include elevated LDL-cholesterol levels in the blood, ■ hypertension, cigarette smoking, diabetes mellitus, physical inactivity, and being overweight or obese.

T E S T YO U R U N D E R S TA N D I N G 1. An open circulatory system (a) is found in flatworms (b) typically includes a hemocoel (c) has a continuous circuit of vessels with openings in the capillaries (d) is characteristic of vertebrates (e) is typically found in animals with a two-chambered heart

10. Norepinephrine (a) slows heart rate (b) is released in cardiac muscle by parasympathetic nerves (c) causes K+ channels in cardiac muscle to open (d) decreases stroke volume (e) causes Ca2+ channels in cardiac muscle to open

2. Which of the following is not a function of the vertebrate circulatory system? (a) helps maintain appropriate pH (b) transports nutrients, oxygen, and metabolic wastes (c) helps maintain fluid balance (d) produces hemocyanin (e) provides internal defense

11. Baroreceptors (a) stimulate renin release (b) activate the renin– angiotensin–aldosterone pathway (c) stimulate sympathetic nerves (d) are stimulated by increased blood pressure (e) send messages to cardiac centers that increase blood pressure

3. Which of the following are most closely associated with oxygen transport? (a) red blood cells (b) platelets (c) neutrophils (d) basophils (e) lymphocytes

12. Label the diagram. See Figure 44-9 to check your answers.

4. Which of the following are most closely associated with blood clotting? (a) red blood cells (b) platelets (c) neutrophils (d) basophils (e) lymphocytes 5. Which choice most accurately describes one sequence of blood flow? 1. right atrium 2. right ventricle 3. left ventricle 4. left atrium 5. pulmonary artery 6. aorta (a) 1, 2, 5 (b) 1, 4, 3 (c) 4, 3, 5 (d) 3, 4, 6 (e) 4, 3, 1 6. Which choice most accurately describes one sequence of blood flow? 1. pulmonary artery 2. pulmonary vein 3. left atrium 4. right atrium 5. pulmonary capillaries (a) 2, 1, 4, 3 (b) 3, 4, 2, 5 (c) 1, 5, 2, 3 (d) 4, 1, 5, 2 (e) 1, 5, 2, 4 7. A cardiac cycle (a) consists of one ventricular heartbeat (b) includes a systole (c) equals stroke volume times heart rate (d) includes a diastole (e) includes a systole and a diastole 8. Blood pressure is determined by (a) cardiac output (b) peripheral resistance (c) blood volume (d) a, b, and c (e) b and c 9. Lymph forms from (a) fluid in the lymph nodes (b) blood serum (c) plasma combined with protein (d) fluid released by lymph nodes (e) interstitial fluid

CRITICAL THINKING 1. When the nerves to the heart are cut, the heart rate increases to about 100 contractions per minute. What does this indicate about the regulation of the heart rate? 2. EVOLUTION LINK. A heart divided into four chambers independently evolved in birds and mammals. What are the advantages of this adaptation? 3. EVOLUTION LINK. How is the heart of a fish specifically adapted to its lifestyle?

4. SCIENCE, TECHNOLOGY, AND SOCIETY. Explain the association between atherosclerosis and ischemic heart disease. Imagine that you were the Surgeon General. What types of programs would you advocate in order to decrease heart disease? As a society, do you think we should invest resources into the development of stem cell therapies for more effectively treating myocardial infarction? Additional questions are available in CengageNOW at www.cengage.com/ login.

45

The Immune System: Internal Defense

Dr. David M. Phillips/Visuals Unlimited/Getty Images

Immune response to infection. This SEM shows a macrophage (large, deep blue cell) engulfing bacteria (small green cells). The SEM has been color-enhanced.

KEY CONCEPTS

T

45.1 During animal evolution, defenses that depend on

organisms and certain toxins. Derived from the Latin for “safe,” the word

protein receptors to recognize dangerous pathogens became part of the genome of every animal.

immune refers to the early observation that when people recovered from

45.2 Nonspecific immune responses provide general and

the same illnesses again.

immediate protection against pathogens, some toxins, and cancer cells.

45.3 Specific immune responses include cell-mediated

he immune system, our internal defense system, is made up of the cells and molecules that protect the body against disease-causing

smallpox and other serious infections, they were safe from contracting Disease-causing organisms, or pathogens, include certain viruses, bacteria, fungi, protozoa, and worms. Pathogens enter the body with air,

immunity and antibody-mediated immunity.

food, and water; during copulation; and through wounds in the skin. The

45.4 In cell-mediated immunity, specific lymphocytes

immune system recognizes pathogens and many types of toxins and rap-

known as T cells are activated; they release proteins that destroy cells infected with viruses or other intracellular pathogens.

idly responds to eliminate them. For example, macrophages act quickly to

45.5 In antibody-mediated immunity, specific lymphocytes known as B cells are activated; they multiply and differentiate into plasma cells, which produce antibodies.

45.6 Some diseases, such as cancer, and disease-causing organisms, for example HIV, the virus that causes AIDS, compromise or overcome the immune system. Some immune responses have negative effects on health: for example, immediate hypersensitivity reactions can result in allergic responses that can cause discomfort, rashes, and even death; and immune responses directed at self can cause autoimmune disease.

destroy invading bacteria (see photograph). The immune system normally functions very effectively. However some pathogens, and also cancer cells, sometimes evade or overcome the body’s internal defenses, resulting in disease. Certain genetic mutations and some pathogens can prevent or compromise immune function. For example, HIV, the retrovirus that causes AIDS, destroys T cells, an important component of the immune system. At times, the immune system overreacts, as in allergic reactions. Immune responses are sometimes directed against our own tissues, resulting in autoimmune disease such as rheumatoid arthritis or multiple sclerosis. At times, the immune system

attacks when we would prefer it did not. For example immune responses can destroy the cells of heart or kidney transplants. Immunology, the study of internal defense systems of humans and other animals, is one of the most rapidly changing and exciting fields of biomedical research. The immune system is a collection of many types of cells and tissues scattered throughout the body. Effective cell signaling among these cells is critical. Cells of the immune system communicate directly by means of their surface molecules and indirectly by releasing messenger molecules. Understanding the complex signaling systems of the immune system is a major focus of research. Investigators are also working to better understand the evolution of the immune system. One of the greatest accomplishments of immunologists has been the development of vaccines that prevent disease. Another has been developing techniques for successful tissue and organ transplantation. Sophisticated research strategies, such as gene transfer, have enabled immunologists to expand their knowledge of the cells and molecules that interact to generate immune re-

that come in contact with an animal’s body. The inflammatory response and phagocytosis are nonspecific defenses that destroy bacteria that manage to penetrate the outer defenses. Specific immune responses are highly specific for distinct macromolecules. Specific immune responses are also referred to as acquired immunity. Any molecule that cells of the immune system specifically recognize as foreign is called an antigen. Proteins are the most powerful antigens, but some polysaccharides and lipids are also antigenic. Specific immune responses are directed toward particular antigens and typically include the production of antibodies, highly specific proteins that recognize and bind to specific antigens. An important characteristic of specific immune responses is immunological memory, which means that the system “remembers” foreign or dangerous molecules and responds more effectively to repeated encounters with the same molecules. Immunologists do not yet understand what signals the immune system not to respond to foreign molecules that are harmless. Some researchers have suggested that the evolution of the immune system allowed animals to manage bacteria and other microbes that are beneficial, as well as to defend against harmful ones.

sponses. These techniques have led to the development of new approaches to the prevention and treatment of disease. Much has been learned, and many challenges lie ahead.

45.1 EVOLUTION OF IMMUNE RESPONSES ■ ■ LEARNING OBJECTIVES 1 2

Distinguish between nonspecific and specific immune responses. Compare, in general terms, the immune responses that have evolved in invertebrates and vertebrates.

An immune response is the process of recognizing foreign or dangerous macromolecules and responding to eliminate them. The immune system distinguishes between self and nonself. Such recognition is possible because each individual is biochemically unique. Cells have surface proteins different from those on the cells of other species or even other individuals of the same species. An animal’s immune system recognizes its own cells and can identify those of other organisms as foreign. Thus, the distinctive macromolecules of a pathogen that invades an animal stimulate the animal’s defensive responses. A single bacterium may have from 10 to more than 1000 distinctive macromolecules on its surface. The immune system also responds to danger signals from injured tissues, such as proteins released when cell membranes are damaged. Two main types of immune responses protect the body: nonspecific and specific (FIG. 45-1). Nonspecific immune responses provide immediate, general protection against pathogens, parasites, some toxins and drugs, and cancer cells. Nonspecific immune responses are referred to as innate immunity. Nonspecific immune responses prevent most pathogens from entering the body and rapidly destroy those that do penetrate the outer defenses. For example, the cuticle or skin provides a physical barrier to pathogens

Invertebrates launch nonspecific immune responses Many invertebrates (cnidarians, annelids, and mollusks) are covered by mucus that traps and kills pathogens. Tough external skeletons, such as cuticles, shells, or chitinous exoskeletons, shield the bodies of nematodes, most mollusks, arthropods, and many other invertebrates (see Chapter 31). All animal species studied have phagocytes that move through the body, engulfing and destroying bacteria and other foreign matter. In mollusks, substances in the hemolymph (blood) enhance phagocytosis. Annelids are the only invertebrates known to have natural killer cells, a type of lymphocyte that destroys tumor cells and cells infected by viruses. Researchers have described more than 800 antimicrobial peptides, chemicals that inactivate or kill pathogens. Investigators have identified antimicrobial peptides in all eukaryotes (including plants) that have been studied, suggesting an early common origin of these molecules. When researchers inject an antigen into an insect, as many as 15 different antimicrobial peptides are produced within a few hours. These chemicals are very effective in part because of their small size (a dozen or fewer amino acids), which facilitates their rapid production and diffusion. What stimulates an animal to produce antimicrobial peptides and other immune defenses? Animal cells have receptors that recognize common types of pathogen molecules and then activate one or more signal transduction pathways, resulting in the production of antimicrobial peptides. All animals studied produce molecules and cells that rush to the location of an infection and produce an inflammatory response. Invertebrates distinguish between their own cells and those of other species. For example, sponge cells have specific glycoproteins on their surfaces that enable them to recognize their own species. When cells of two different species are mixed, they sort themselves out according to species. When two different species

PATHOGENS INVADE TISSUES

Rapid

Requires several hours to several days

NONSPECIFIC (INNATE) IMMUNE RESPONSES

SPECIFIC (ADAPTIVE) IMMUNE RESPONSES

Inflammation

Barriers, for example, epithelial lining of digestive and respiratory systems

Pattern recognition receptors

Cytokines and complement

NK cells

Dendritic cells

Phagocytes (macrophage shown)

Cellmediated immunity

Antibodymediated immunity

Phagocytosis

Antimicrobial peptides and other compounds

FIGURE 45-1 Animated Overview of human immune responses Nonspecific immune responses include barriers that prevent the entrance of pathogens; pattern recognition receptors; cytokines and complement; natural killer (NK) cells, dendritic cells, and phagocytes; and inflammation. During inflammation, certain cells release cytokines that activate specific immune responses: cell-mediated immunity and antibody-mediated immunity.

of sponges are forced to grow in contact with each other, tissue is destroyed along the region of contact. Cnidarians (such as corals), annelids (such as earthworms), arthropods (such as insects), and echinoderms (such as sea stars) reject tissue grafted from other animals, even from the same species. Certain cnidarians, arthropods, some echinoderms, and simple chordates (such as tunicates) appear to remember antigens for a short period. As mentioned earlier, immunological memory enables the body to respond more effectively when it encounters the same pathogens again. Although certain invertebrates demonstrate some specificity and memory, their immune responses are primarily nonspecific. Echinoderms and tunicates are the most ancient groups of animals known to have differentiated white blood cells that perform limited immune functions. Investigators have identified two genes in the purple sea urchin that are very similar to vertebrate genes

important in producing antibodies. This discovery suggests that some invertebrates may have some capacity to launch specific immune responses.

Vertebrates launch nonspecific and specific immune responses Like invertebrates, vertebrates depend on nonspecific immune responses for their initial assault on invading pathogens. However, their specific immune responses rapidly fine-tune the more general nonspecific responses. The specialized lymphatic system that has evolved in vertebrates includes cells such as lymphocytes, white blood cells specialized to carry out immune responses, and organs such as lymph nodes (see Chapter 44). Specific immune responses, including action by lymphocytes and antibodies, may have first evolved in

jawed vertebrates. In the discussion that follows, we focus on the human immune system, with references to immune mechanisms of other animals.

Review ■

■

What are two key ways in which specific immune responses differ from nonspecific immune responses? In general, how do vertebrate immune responses differ from invertebrate responses?

45.2 NONSPECIFIC IMMUNE RESPONSES ■ ■ LEARNING OBJECTIVE 3

Describe nonspecific immune responses, including physical barriers; recognition by pattern recognition receptors; actions of phagocytes and other cells; actions of cytokines and complement; and the inflammatory response.

Nonspecific immune responses are also referred to as innate immunity. Nonspecific immune mechanisms are always present and ready to respond rapidly to invading pathogens. Nonspecific immune responses provide immediate, general protection against pathogens, parasites, some toxins and drugs, and cancer cells. Nonspecific defenses include physical barriers, such as the skin; action by pattern recognition receptors; action of proteins such as cytokines and those that make up the complement system; actions by cells, including phagocytes, dendritic cells, and natural killer (NK) cells; and the inflammatory response.

Physical barriers prevent most pathogens from entering the body An animal’s first line of defense is its outer covering—cuticle, shell, chitin, or skin—which blocks the entry of pathogens. The digestive (gastrointestinal) tract, respiratory passageways, urinary tract, and vagina all open to the outside of the body. The epithelial linings of these passageways are barriers between the outside environment and the inside of the body. Various mechanisms within epithelial barriers expel, trap, or destroy pathogens. For example, pathogens that enter the body with inhaled air may be filtered out by hairs in the nose or trapped in the sticky mucous lining of the respiratory tract. Once trapped, they are destroyed by phagocytes. Physical barriers are also equipped with chemical weapons. For example, antimicrobial peptides called defensins are produced by the epithelium of the skin of many animals, including mammals. Defensins, which are also found in saliva, are toxic to bacteria and fungi. Mucus contains a substance called mucin, which chemically destroys invaders. Lysozyme, an enzyme found in tears and other body fluids, attacks the cell walls of many gram-positive bacteria. Mucus, tears, urine, and other body fluids flush pathogens out of the body. Invading bacteria must also compete with the populations of microbes that inhabit our skin and other surface tissues. These resident microbes are normally not harmful.

Pattern recognition receptors activate nonspecific immune responses When pathogens breach the body’s outer barriers, their chemical properties activate other nonspecific defense responses, and most invaders are quickly eliminated. Bacteria and other microbes have certain proteins and other molecules that are not present on the cells of humans and other mammals. These chemical compounds (or specific parts of molecules) that stimulate nonspecific immune responses are called pathogen-associated molecular patterns, or PAMPs. Each group of microbes—viruses, gram-positive bacteria, and fungi—share certain PAMPs. For example, certain viruses have double-stranded RNA and gram-positive bacteria have peptidoglycan. Pattern recognition receptors have evolved that recognize PAMPs as nonself. When these receptors on phagocytes bind with PAMPs, nonspecific immune responses are activated. For example, the phagocytes become more effective at taking in and destroying the pathogens. These receptors may also transduce signals that stimulate the inflammatory response. The most important group of pattern recognition receptors, the Toll-like receptors, are expressed on several types of cells. Toll-like receptors (homologous to a protein known as the Toll protein in Drosophila) activate certain transcription factors. These transcription factors stimulate the expression of genes that encode certain proteins, such as cytokines, which are important signaling molecules.

Cells of the nonspecific immune system destroy pathogens and produce chemicals Nonspecific immune responses depend on the action of several types of cells, including phagocytes, natural killer cells, and dendritic cells. Neutrophils, the most common type of white blood cell (see Chapter 44), and macrophages are the main phagocytes in the body. Recall that phagocytosis is the process by which cells engulf microorganisms, foreign matter, or other cells (see Fig. 5-21). A neutrophil can phagocytose about 20 bacteria before it becomes inactivated (perhaps by leaking lysosomal enzymes) and dies. Macrophages are large phagocytes that develop from monocytes, a type of nongranular white blood cell. A macrophage can phagocytose about 100 bacteria during its lifespan (FIG. 45-2). Macrophages also release antiviral agents. Some macrophages patrol the body’s tissues, engulfing damaged cells or foreign matter (including bacteria). Other macrophages stay in one place and destroy bacteria that pass by. For example, air sacs in the lungs contain large numbers of macrophages that destroy foreign matter entering with inhaled air. Vertebrate macrophages have Toll-like receptors that recognize certain PAMPs and respond by producing cytokines. For example, a lipopolysaccharide found on gram-negative bacteria stimulates macrophages. In response, macrophages release cytokines that enhance the inflammatory response. Can bacteria counteract the phagocyte’s attack? Many strategies have evolved in bacteria that protect them from their hosts. Streptococcus pneumoniae, for example, has cell walls or capsules that resist the action of the phagocyte’s lysosomal enzymes. Other

Lennart Nilsson, Boehringer Ingelheim International GmbH

Lennart Nilsson, Boehringer Ingelheim International GmbH

5 μm

Lennart Nilsson, Boehringer Ingelheim International GmbH

(a) A macrophage extends a pseudopod toward an invading Escherichia coli bacterium that is already multiplying.

0.5 μm

(b) A bacterium is trapped within the engulfing pseudopod.

FIGURE 45-2 Phagocytosis Macrophages are efficient warriors. These colorized SEMs show macrophages phagocytosing bacteria.

bacteria release enzymes that destroy lysosomal membranes. The powerful lysosomal enzymes then spill out into the cytoplasm and may destroy the phagocyte. Natural killer (NK) cells are large, granular lymphocytes that originate in the bone marrow. They account for about 10% of circulating lymphocytes. NK cells are active against tumor cells and cells infected with some types of viruses. They destroy target cells by both nonspecific and specific (antibody-mediated) processes.

5 μm

(c) A macrophage takes in trapped bacteria along with its own plasma membrane. The macrophage plasma membrane will seal over the bacteria, and powerful lysosomal enzymes will destroy them.

NK cells release cytokines, as well as enzymes that destroy target cells. Perforins are enzymes that cause pores to form in the plasma membrane of the target cell, allowing granzymes, another type of enzyme, to enter the cell. Granzymes then activate a cascade of reactions that cause the cell to self-destruct by apoptosis, programmed cell death. Several cytokines stimulate NK cell activity. When NK cell levels are high, resistance to certain cancers may be strengthened. Psychological stressors may decrease NK cell activity and thus enhance tumor growth. Biologists first identified dendritic cells in 1868 and then rediscovered them in 1973, but the small numbers of these cells made them hard to study until sophisticated techniques became available in the 1990s. Their name derives from their long cytoplasmic extensions called dendrites (not to be confused with the dendrites of neurons). Dendritic cells develop from precursor cells in the bone marrow and from monocytes. Dendritic cells are strategically stationed in all tissues of the body that come into contact with the environment—the skin and the epithelial linings of the digestive, respiratory, urinary, and vaginal passageways. When pathogens infect a tissue, PAMPs activate dendritic cells. One type of dendritic cell responds to viral infection. Dendritic cells produce interferons, cytokines that are antiviral. Certain types of activated dendritic cells capture microbial antigens by phagocytosis or by receptor-mediated endocytosis (see Chapter 5).

Cytokines and complement mediate immune responses Cells of the immune system secrete a remarkable number of regulatory and antimicrobial peptides and proteins. Two major groups

of molecules that are critical in nonspecific defense responses are cytokines and complement. Cytokines are important signaling molecules Cytokines are a large, diverse group of peptides and proteins (most are glycoproteins) that serve as important signaling molecules and perform regulatory functions. The same kinds of cytokines can be produced by many different types of cells. Also, a cytokine may affect many different kinds of cells. Just as the functions of nonspecific and specific immune responses overlap, the actions of cytokines of these subsystems also overlap. For example, cytokines produced by nonspecific cells such as macrophages can activate lymphocytes involved in specific immune responses. Cytokines help regulate the intensity and duration of immune responses. They are also important in regulating many other biological processes, such as cell growth, repair, and cell activation. Cytokines bind to specific membrane receptors on target cells. These signaling molecules can act as autocrine agents, which affect the very cells that produce them, or as paracrine agents, which regulate the activity of nearby cells (see Chapter 49). When produced in large amounts, some cytokines act as hormones that circulate in the blood and affect distant tissues. Biologists name the types of cytokines for their function or origin. We discuss several examples, including interferons, tumor necrosis factors, interleukins, and chemokines. When infected by viruses or other intracellular parasites (some types of bacteria, fungi, and protozoa), certain cells respond by secreting cytokines called interferons. Type I interferons are produced by either macrophages or fibroblasts, cells that produce the fibers of connective tissues. Type I interferons inhibit viral replication and activate NK cells that have antiviral actions. Viruses produced in cells exposed to Type I interferons are not very effective at infecting other cells. Interestingly, even though Type I interferons are produced as part of the nonspecific immune response, once released they can regulate the action of certain cells of the specific immune response. This interaction of nonspecific and specific immune mechanisms is an example of the power and complexity of the immune system. Type II interferon (also called IFN-gamma) is critical in both nonspecific and specific immunity. In nonspecific immunity, Type II interferon stimulates macrophages to destroy tumor cells and host cells that have been infected by viruses. (The interferon activates transcription of genes that encode enzymes needed to produce antimicrobial compounds in lysosomes.) Since their discovery in 1957, interferons have been the focus of a great deal of research. Pharmaceutical companies now use recombinant DNA techniques to produce large quantities of some interferons. The U.S. Food and Drug Administration (FDA) has approved interferons for treating several diseases, including hepatitis B and hepatitis C, genital warts, several types of cancer (for example, leukemia), a type of multiple sclerosis, and AIDS-related Kaposi’s sarcoma. In response to gram-negative bacteria, certain other pathogens, or some tumors, tumor necrosis factor (TNF) stimulates immune cells to initiate an inflammatory response. In fact, TNF was so-named because it was first identified as a compound that destroyed tumor cells. TNF is secreted mainly by macrophages and

by T cells, a type of lymphocyte. In severe infections, large amounts of TNF are produced. TNF is then circulated in the blood and can have widespread effects. For example, it acts on the hypothalamus, inducing fever. Sometimes infection by gram-negative bacteria, such as Salmonella typhi, results in the release of large amounts of TNF and other cytokines. This results in a cascade of reactions leading to septic shock, a potentially fatal condition that may involve high fever and malfunction of the circulatory system. Thus, cytokines can sometimes have harmful effects. Interleukins are a diverse group of cytokines secreted mainly by macrophages and lymphocytes. Interleukins are numbered according to their order of discovery. They regulate interactions between white blood cells, such as lymphocytes and macrophages, and other cells. Interleukin-1 (IL-1) functions with TNF in mediating inflammation. Interleukin-12 (IL-12) stimulates NK cells, as well as T cells, to produce IFN-gamma. Some interleukins have widespread effects. During infection IL-1 can reset the body’s thermostat in the hypothalamus, resulting in fever and its symptoms. The overproduction of certain interleukins has been associated with clinical disease, including atherosclerosis. Chemokines, a large group of cytokines, are signaling molecules that attract, activate, and direct the movement of various cells of the immune system. For example, they regulate the migration of white blood cells from the blood to the tissues. Some chemokines are produced in response to infection and are mediators of the inflammatory response. Complement promotes destruction of pathogens and enhances inflammation Phagocytes secrete cytokines that activate the complement system. Complement, so named because it “complements” the action of other defensive responses, consists of more than 20 proteins present in plasma and other body fluids. Similarities in complement proteins in many species, including horseshoe crabs, sea urchins, tunicates, and mammals, suggest that these molecules evolved millions of years ago and have been highly conserved. Normally, complement proteins are inactive until the body is exposed to an antigen. Certain pathogens activate the complement system directly. In other cases, the binding of an antigen and antibody stimulates activation. Complement activation involves a cascade of reactions; each component acts on the next in the series. Proteins of the complement system then work to destroy pathogens. Complement proteins are active against many antigens, and their actions are nonspecific: they (1) lyse viruses, bacteria, and other cells; (2) coat pathogens, making them less “slippery” so that macrophages and neutrophils phagocytose them more easily; (3) attract white blood cells to the site of infection, a process called chemotaxis; and (4) bind to specific receptors on cells of the immune system, stimulating specific actions, such as secreting regulatory molecules and enhancing the inflammatory response.

Inflammation is a protective response The inflammatory response, or inflammation, is one of the body’s key responses to infection or injury. Tissue injury caused by patho-

gen invasion or physical injury activates a clotting factor in the blood plasma that turns on three different interconnecting molecular cascades in the plasma, including the clotting cascade. Many of these reactions generate molecules—for example, the peptide bradykinin—that mediate the inflammatory process. These plasma mediators dilate blood vessels and increase capillary permeability (FIG. 45-3 and 45-4). Many cytokines signal white blood cells to launch the inflammatory response, and some cytokines help regulate the process. Inflammation also activates the complement system. The clinical characteristics of inflammation are heat, redness, edema (swelling), and pain. The inflammatory response includes three main processes.

Fluid and antibodies leave the circulation and enter the tissues. As the volume of interstitial fluid increases, edema occurs. The edema, along with the action of certain enzymes in the plasma, causes the pain that characterizes inflammation. 3. Increased phagocytosis. Increased blood flow brings large numbers of neutrophils and other phagocytic cells to the inflamed region within a few hours of tissue injury or infection. The phagocytes migrate out of the capillaries and into the infected tissues. Macrophages secrete chemokines and other cytokines that recruit and activate additional neutrophils. One of the main functions of inflammation is increased phagocytosis.

1. Vasodilation. Macrophages and mast cells stationed in the tissues respond rapidly to damaged tissue or infection. Mast cells, which are found in most tissues close to blood vessels, release histamine, cytokines, and other compounds that dilate blood vessels in the affected area. Expanded blood vessel diameter brings more blood to the area. The increased blood flow warms the skin and reddens skin that contains little pigment. 2. Increased capillary permeability. Histamine and other compounds released by mast cells increase capillary permeability.

Although the inflammatory response begins as a local response, sometimes the entire body may become involved. Fever, a common clinical symptom of widespread inflammation, helps the body fight infection. Elevated body temperature increases phagocytosis and interferes with the growth and replication of microorganisms. Fever breaks down lysosomes, destroying cells infected by viruses. Fever also promotes the activity of certain lymphocytes. A shortterm, low fever speeds recovery. The inflammatory response is vital in overcoming infection, but chronic inflammation contributes to many chronic diseases.

KEY POINT

During the inflammatory response, capillaries dilate and become more permeable, bringing more phagocytes to the infected area; phagocytosis increases.

PATHOGENS INVADE TISSUES

Cells injured

Activate molecules in plasma, mast cells, macrophages

Vasodilation

Increased capillary permeability

Increased blood flow to area

Brings phagocytes, nutrients, antibodies

Redness

Antibodies and phagocytes pass from blood into inflamed area

Phagocytes migrate to region

Edema

Increased temperature

Pain

Release TNF, IL-1

Fever

FIGURE 45-3 Animated Overview of the inflammatory response When skin is injured, bacteria (and other pathogens) enter the tissues. Tissue injury activates a clotting factor in the blood that turns on three molecular pathways in the plasma. Some of the molecules produced dilate blood vessels and increase capillary permeability.

Increased phagocytosis

Systemic response

Capillary

1

Bacteria invade tissue. Injury activates clotting factors in blood. These factors turn on pathways that produce molecules that dilate blood vessels and increase permeability.

1

Bacteria Cytokines 2

2

Activated macrophages and neutrophils phagocytose bacteria. Macrophages secrete cytokines.

3

Activated mast cells release histamine and other compounds.

Macrophage

3

Neutrophil

Histamine

Mast cells

(a) Histamine, cytokines, and other compounds released by mast cells increase vasodilation and capillary permeability. Certain cytokines attract phagocytic cells. Macrophages and neutrophils in the infected area engulf and destroy bacteria. Macrophages secrete cytokines that recruit additional neutrophils. Mast cells secrete histamine, cytokines, and other substances that dilate blood vessels.

FIGURE 45-4 Animated The inflammatory response Bacteria are represented by brown rods; cytokines by green dots; histamines by orange squares; and other mast cell secretions by orange dots.

For example, inflammation has been linked with atherosclerosis, rheumatoid arthritis, Crohn’s disease, diabetes, Alzheimer’s disease, and cancer. Although we do not yet understand the details, chronic inflammation appears to occur when homeostatic mechanisms that normally turn off the inflammatory response do not function appropriately. Investigators have identified prolonged infections, cigarette smoking, gum disease, and obesity as risk factors for chronic inflammation.

Review ■ ■ ■ ■

■

What barriers keep most pathogens from invading the body? What is the function of pattern recognition receptors? What is the function of NK cells? macrophages? What are the main groups of cytokines? What is the function of each group? What processes are involved in the inflammatory response?

45.3 SPECIFIC IMMUNE RESPONSES ■ ■ LEARNING OBJECTIVES 4 Contrast cell-mediated and antibody-mediated immunity, and give an 5

overview of each process. Describe the functions of the principal cells of the specific immune system and of the major histocompatibility complex.

Specific immune responses are also referred to as adaptive immunity or acquired immunity. In contrast to nonspecific immune mechanisms that are always present and ready to respond to infection, specific immune responses develop in response to an infection and adapt to the specific pathogens invading the body. These immune responses are highly specific for distinct macromolecules.

Capillary dilated and permeable

4

Cytokines 4

Histamine and cytokines dilate blood vessels and increase capillary permeability.

5

Cytokines attract phagocytes to the infection site.

6

Increased phagocytosis.

Neutrophil

5

Histamine Mast cells

Monocyte

Macrophage

6

Neutrophils

(b) Large numbers of monocytes and phagocytic cells pass through the now dilated, permeable capillaries into the infected area. Monocytes develop into macrophages. Phagocytosis increases.

While nonspecific immune responses destroy pathogens and prevent the spread of infection, the body mobilizes its specific immune responses. It takes several days to activate specific immune responses, but once in gear, these mechanisms are extremely effective. As discussed earlier, specific immune responses are precisely targeted to destroy specific antigens, and they have immunological memory. Two types of specific immune responses are cell-mediated immunity and antibody-mediated immunity.

Many types of cells are involved in specific immune responses The main types of cells of the specific immune system are shown in FIGURE 45-5. Two key cell types that participate in specific immune responses are lymphocytes and antigen-presenting cells. Phagocytes and many other cells important in nonspecific immune responses also participate in specific responses. Eosinophils, for example, are white blood cells that release proteins, including cytokines, from their granules. Some of these cytokines are important in spe-

cific immune responses. Eosinophils also release proteins that are toxic to parasitic worms. Lymphocytes are the principal warriors in specific immune responses Lymphocytes are important in both specific and nonspecific immunity. Natural killer cells (NK cells) were discussed in the previous discussion of nonspecific immunity. Recall that NK cells are lymphocytes that kill virally infected cells and tumor cells. T lymphocytes, or T cells, and B lymphocytes, or B cells, function in specific immunity. T cells are responsible for cell-mediated immunity. They are the body’s cellular soldiers. T cells travel to the site of infection and attack body cells infected by invading pathogens, as well as foreign cells (such as those introduced in tissue grafts or organ transplants). T cells also destroy cells altered by mutation (cancer cells). B cells are responsible for antibody-mediated immunity. B cells mature into plasma cells, which produce specific antibodies. Antibodies bind to specific antigens, neutralizing them or marking

KEY POINT

Cells important in specific immune responses include antigen-presenting cells (macrophages and dendritic cells) and lymphocytes (T cells and B cells). These cells interact through complex signaling.

Bone marrow stem cells

B cells mature in bone marrow

T cells mature in thymus

Monocyte

Macrophage

Dendritic cell

n T cell Antige tion ta presen n n tige atio An sent Migrates to lymph nodes pre

Memory T cell

T cytotoxic cell

T helper cell

Cell-mediated immunity

B cell

n tio ra e op Co

Migrates to lymph nodes

Plasma cell

Memory B cell

Antibody-mediated immunity

FIGURE 45-5 Some key cells in specific immune responses

them for destruction. Antibody-mediated immunity is one of the body’s chief chemical defense strategies. All lymphocytes develop from stem cells in the bone marrow. B cells complete their development in the adult bone marrow. T cells mature in the thymus gland (from which the T in their name derives). Large numbers of mature lymphocytes reside in lymph organs, including the spleen, lymph nodes, tonsils, and other lymph tissues strategically positioned throughout the body. Although T cells and B cells are similar in appearance when viewed with a light microscope, sophisticated techniques such as fluorescence microscopy demonstrate that these cells can be differentiated by their unique cell-surface macromolecules. T cells and B cells have different functions and life histories and tend to locate in (or “home” to) separate regions of the spleen, lymph nodes, and other lymph tissues.

Millions of B cells are produced in the bone marrow daily. Each B cell is genetically programmed to encode a glycoprotein receptor that can bind with a specific type of antigen. When a B cell comes into contact with an antigen that binds to its receptors, the B cell becomes activated. B-cell activation is a complex process that typically requires the participation of a particular type of T cell. Once activated, a B cell divides rapidly to form a clone of identical cells. The cloned B cells differentiate into plasma cells, which produce antibodies. A plasma cell can produce more than ten million molecules of antibody per hour! The antibody binds to the antigen that originally activated the B cells. Some activated B cells do not become plasma cells but instead become long-living memory B cells that continue to produce small amounts of antibody after the body overcomes an infection.

On their way from the bone marrow to the lymph tissues, T cells are processed in the thymus gland. The thymus makes T cells immunocompetent, capable of immunological response. As T cells move through the thymus, they divide many times and develop specific surface proteins with distinctive receptor sites. The T-cell population undergoes positive and negative selection. In negative selection, T cells in the thymus that react to selfantigens undergo apoptosis. Immunologists estimate that more than 90% of developing T cells are negatively selected. In positive selection, T cells are allowed to mature if they recognize self-antigens and bind with foreign antigens. These T cells differentiate and leave the thymus to take up residence in other lymph tissues or to launch immune responses in infected tissues. By selecting only appropriate T cells, the thymus gland ensures that T cells can distinguish between the body’s own molecules and foreign antigens. Most T cells in the thymus differentiate just before birth and during the first few months of postnatal life. If the thymus is removed before this processing takes place, an animal does not develop cell-mediated immunity. If the thymus is removed after that time, cell-mediated immunity is less seriously impaired. T cells are distinguished by the T-cell receptor (TCR), which recognizes specific antigens. Two main populations of T cells are T cytotoxic cells and T helper cells. T cytotoxic (TC) cells are also known as CD8 cells because they have a glycoprotein designated CD8 on their plasma membrane surface. Known less formally as killer T cells, TC cells recognize and destroy cells with foreign antigens on their surfaces. Their target cells include virus-infected cells, cancer cells, and foreign tissue grafts. T helper (TH) cells, also known as CD4 cells, have a surface glycoprotein designated CD4. TH cells secrete cytokines that activate B cells, T cells, and macrophages. Regulatory T cells, known as Tregs, are a subpopulation of TH cells. Tregs help regulate immune responses by suppressing the function of certain T cells. After an infection, memory T cells (both memory TC cells and memory TH cells) remain in the body. Antigen-presenting cells activate T helper cells Macrophages, dendritic cells, and B cells function as “professional” antigen-presenting cells (APCs) that display foreign antigens as well as their own surface proteins. APCs are inactive until their pattern recognition receptors recognize PAMPs on pathogens. Once activated, an APC ingests the pathogen. Lysosomal enzymes inside the APC degrade most, but not all, of the bacterial antigens. The APC displays fragments of the foreign antigens on its cell surface in association with a type of self-molecule (class II MHC, discussed later). The activated APC expresses additional signaling molecules, called costimulatory molecules, along with the displayed antigen. These fully functional APCs present displayed antigens to T cells. In addition to their function as APCs, macrophages secrete more than 100 different compounds, including cytokines and enzymes that destroy bacteria. When stimulated by bacteria, macrophages secrete interleukins that activate B cells and certain T cells. We have seen that interleukins also promote nonspecific immune responses, such as fever. Receptors on dendritic cells bind antigens of microbes. Dendritic cells take these antigens, process them, and display antigen

fragments on their cell surface. Now APCs, the dendritic cells leave the epithelial lining and, guided by chemokines, migrate to the lymph nodes. Costimulatory molecules, along with the displayed antigen, attract and activate specific T cells capable of responding to the antigen. Thus, dendritic cells are specialized to bind, process, transport, and present antigens to T cells.

The major histocompatibility complex is responsible for recognition of self The ability of the vertebrate immune system to distinguish self from nonself depends largely on a cluster of closely linked genes known as the major histocompatibility complex (MHC). In humans, the MHC is also called the human leukocyte antigen complex (HLA). The MHC genes are polymorphic (variable). Within the population there are multiple alleles, more than 40 for each locus. As a result, the cell-surface proteins for which they code are generally different in each individual. With so many possible combinations, no two people, except identical twins, are likely to have all the same MHC proteins on their cells. The more closely related two individuals are, the more MHC genes they have in common. You can think of your MHC proteins as a biochemical “fingerprint.” The MHC genes encode MHC antigens, referred to as selfantigens, which differ in chemical structure, function, and tissue distribution. Class I MHC genes encode glycoproteins expressed on the surface of most nucleated cells. These antigens bind with foreign antigens from viruses or other pathogens within the cell to form a foreign antigen–class I MHC glycoprotein complex. The cell displays this complex on its surface and presents it to TC cells. Thus, any infected cell can function as an antigen-presenting cell and can activate certain TC cells. Class II MHC genes encode glycoproteins expressed primarily on “professional” APCs—dendritic cells, macrophages, and B cells. These class II MHC antigens combine with foreign antigen from bacteria, and the cell presents the complex to TH cells. Class III MHC genes encode many secreted proteins involved in the immune response, including components of the complement system and TNFs.

Review ■

■ ■

How are cell-mediated immunity and antibody-mediated immunity different? What are two main types of T cells? What is the function of B cells? What is the function of antigen-presenting cells? Describe two types.

45.4 CELL-MEDIATED IMMUNITY ■ ■ LEARNING OBJECTIVE 6 Describe the sequence of events in cell-mediated immunity.

T cells and APCs (mainly dendritic cells and macrophages) are responsible for cell-mediated immunity. T cells destroy cells infected with viruses and macrophages that have ingested “smart” bacteria. These bacteria escape from the vesicles in which they are enclosed during phagocytosis. They escape destruction by lysosomal

KEY POINT

T cells and APCs are responsible for cell-mediated immunity. Ingested pathogen

T-cell receptor APC

1

TH cell is activated by specific foreign antigen–MHC complex presented by APC.

Antigen–MHC complex

TH cell

2

Activated TH cell increases in size and divides by mitosis.

3

Clonal expansion: clone of competent TH cells produced.

Memory T cells 4

TH cells differentiate and migrate to infected tissues; they release cytokines that act on other T cells and on macrophages.

5

TC cells are activated by foreign antigen–MHC complex displayed by infected cells. TC cells form clones (not shown) and migrate to infected tissues.

6

TC cells release proteins that destroy infected cells.

Cytokines TH cell TC cells Antigen–MHC complex Infected cell

Infected cell destroyed

FIGURE 45-6 Animated Cell-mediated immunity When activated by a foreign antigen–MHC complex presented by an APC and by cytokines, a TH cell divides and gives rise to a clone of TH cells. TC cells are also activated by antigens presented on the surfaces of infected cells, and they also form a clone. Both types of T cells migrate to the site of infection. TH cells release cytokines that stimulate TC cells and activate macrophages so that they phagocytose pathogens. TC cells release proteins that destroy infected cells.

enzymes by “hiding” in the cytoplasm of macrophages. T cells destroy cells that have been altered in some way, such as cancer cells. They also destroy the cells of foreign grafts such as a transplanted kidney or liver. How do T cells know which cells to target? T cells do not recognize antigens unless they are presented properly. Recall that when a virus (or other pathogen) infects a cell, some of the viral protein is broken down to peptides and displayed with class I MHC molecules on the cell surface. Only T cells with receptors that bind to the specific antigen– MHC complex become activated. There are two general populations of T cells: TH cells and TC cells. Each TH cell expresses the glycoprotein called CD4 and each TC cell expresses CD8. Within each general population of T cell, there are thousands of different antigen specificities that the T-cell receptors (TCRs) can recognize. Activation of TH cells, for example, occurs when phagocytic cells digest antigen and parts of the antigen are placed within their surface MHC complexes. They then present these antigens to the TCR on TH cells as part of a foreign antigen– class II MHC glycoprotein complex. Once activated by the foreign antigen–MHC class II antigen complex displayed on the surface of APCs, the activated TH cells secrete interleukin 2. This cytokine is a growth factor that stimulates the activated cells to proliferate. The TH cells divide several times, giving rise to a clone of TH cells (FIG. 45-6). This process is called clonal expansion. Some of the TH cells produced migrate to the site of infection. These cells can produce many different types of cytokines. Some attract macrophages to the site of infection and promote the destruction of intracellular pathogens. Others function mainly in promoting antibody-mediated immunity. Each TC cell expresses the CD8 molecule on its surface, as well as more than 50,000 identical TCRs that bind to one specific type of antigen. TC cells recognize antigens presented to them as part of a foreign antigen–class I MHC glycoprotein complex. Generally, fewer than 1 in 100,000 T cells have the same antigen specificity and can respond. After clonal expansion, as many as 1 in 10 TC cells may be specific for the targeted antigen. TC cell activation requires at least two signals in addition to the presented antigen: a costimulatory signal and an interleukin signal. Once activated, a TC cell increases in size and gives rise to a clone of many effector TC cells. Effector TC cells make up the cell infantry. They leave the lymph nodes and make their way to the infected area, where they destroy target cells within seconds after contact. When a TC cell combines with antigen on the surface of the target cell, it destroys the cell in much the same way that NK cells destroy their target cells. The TC secretes perforins that perforate the plasma membrane of the target cell. Granzymes then induce the cell to self-destruct by apoptosis. After releasing cytotoxic substances, the T cell disengages itself from its victim cell and seeks out a new target. The sequence of events for production and function of TC cells in cell-mediated immunity can be summarized as follows:

virus invades body cell ¡ infected cell displays foreign antigen– MHC class I antigen complex ¡ specific TC cell activated by this complex ¡ clone of TC cells produced ¡ effector TC cells migrate to area of infection ¡ TC cells stimulated by cytokines released by TH cells ¡ TC cells release enzymes that destroy

Bacterium Antigen

Antigen

Bacterium

target cells APC

Review ■ ■ ■

How do TH cells function in cell-mediated immunity? How do APCs function in cell-mediated immunity? How do TC cells function in cell-mediated immunity?

B cell Antigen– MHC complex 1a

45.5 ANTIBODY-MEDIATED IMMUNITY

APC takes in bacterium, breaks it down, and presents foreign antigens on its surface in combination with MHC class II antigens.

■ ■ LEARNING OBJECTIVES

1b

B cell independently combines with foreign antigens on surface of bacterium, but B cell is typically not active until stimulated by TH cells.

Cytokines

7 Summarize the sequence of events in antibody-mediated immunity,

APC

including the effects of antigen–antibody complexes on pathogens.

8 Describe the basic structure and function of an antibody, and explain the

TH cell

basis of antibody diversity. 9 Describe the basis of immunological memory, and contrast secondary and primary immune responses. 10 Compare active and passive immunity, and give examples of each.

We have discussed two types of molecules that recognize specific antigens: T-cell receptors and MHC molecules. Here we turn our attention to the third type, antibodies. B cells are responsible for antibody-mediated immunity. A given B cell can produce many copies of one specific antibody. Antibody molecules serve as cellsurface receptors that combine with antigens. Only a B cell with a receptor complementary in shape to a particular antigen can bind that antigen. The antigen enters the B cell by receptor-mediated endocytosis (see Chapter 5). The antigen is then degraded into peptide fragments, and the B cell displays peptide fragments together with MHC protein class II on its surface. In most cases, the activation of B cells involves APCs and TH cells (FIG. 45-7). TH cells stimulate B cells to divide and produce antibodies. However, the TH cell itself must first be activated. TH cells do not recognize an antigen that is presented alone. The antigen must be presented as part of a foreign antigen–class II MHC complex on the surface of an APC. When an APC displaying a foreign antigen–MHC complex contacts a TH cell with complementary T-cell receptors, a complicated interaction occurs. Multiple chemical signals are sent back and forth between cells. For example, the APC secretes interleukins, such as IL-1, that activate TH cells. B cells serve as APCs to T cells. An activated TH cell binds with the foreign antigen–MHC complex on the B cell. The TH cell then releases interleukins, which together with antigen activate the B cell. Once activated, a B cell divides by mitosis, giving rise to a clone of identical cells (FIG. 45-8). This clonal expansion in response to a specific antigen is known as clonal selection (discussed later). The cloned B cells mature into plasma cells that secrete antibod-

Receptor

2 TH

cells are activated when their receptors combine with foreign antigen–MHC antigen complexes on APC. Signaling takes place via cytokines secreted by both cells. TH cell

Foreign antigen

Activated B cell

Cytokines

Cell division

3

Activated TH cell recognizes antigen–MHC complex on B cell. TH cell secretes cytokines that activate B cell that has interacted with foreign antigen. B cell then divides, and subsequent divisions produce clone of identical B cells.

FIGURE 45-7 Animated B-cell activation This figure is a detailed illustration of the first four steps in Figure 45-8.

ies. These antibodies are specific to the antigen that activated the original B cell. It is important to remember that the specificity of the clone is determined before the B cell encounters the antigen. Unlike T cells, most plasma cells do not leave the lymph nodes. Only the antibodies they secrete pass out of the lymph tissues and make their way via the lymph and blood to the infected area. The

KEY POINT

B cells, APCs, and TH cells are responsible for antibody-mediated immunity.

APC 1

APC presents foreign antigen–MHC complex on its surface.

2

TH cell binds with complex and secretes cytokines.

3

Cytokines and antigens activate B cell with receptor that matches antigen.

4

Activated B cell increases in size and divides by mitosis.

Antigen

TH cell

Cytokines

B cell

Activated B cell

Bacterium

B cells

Plasma cells Antibody

5

Clonal expansion: clone of competent B cells is produced.

6

B cells differentiate into plasma cells and memory B cells.

Memory B cells

7

Plasma cells secrete specific antibodies, which are transported via lymph and blood to infected region.

8

Antibodies combine with antigens on pathogen surface, forming antigen–antibody complexes. Pathogens destroyed.

Bacteria with antigen

Antigen–antibody complex

FIGURE 45-8 Animated Antibody-mediated immunity A specific B cell may be activated when a specific antigen binds to immunoglobulin receptors on the B-cell surface. Typically, the B cell also requires cytokine stimulation from an activated TH cell (see Fig. 45-7). Once activated, the B cell divides and produces a clone of B cells. These

cells differentiate and become plasma cells that secrete antibodies. The plasma cells remain in the lymph tissues, but the antibodies are transported to the site of infection by the blood or lymph. After the infection has been overcome, memory B cells remain in the tissues.

secreted antibody is a soluble form of the glycoprotein receptor of the activated B cell. The sequence of events in antibody-mediated immunity can be summarized as follows: pathogen invades body ¡ APC phagocytoses pathogen ¡ foreign antigen–MHC complex displayed on APC surface ¡ TH cell binds with foreign antigen–MHC complex ¡ activated TH cell interacts with a B cell that displays the same antigen ¡ B cell activated ¡ clone of B cells produced ¡ B cells differentiate, becoming plasma cells ¡ plasma cells secrete antibodies ¡ antibodies form complexes with pathogen ¡ pathogen is destroyed

Antigenic determinants

Binding sites Variable (V) region

Light chain

Constant (C) region

Heavy chain Antibody

Antigen

Disulfide bond

(a) The antibody molecule is composed of two light chains and two heavy chains joined by disulfide bonds. The constant (C) and variable (V) regions of the chains are labeled. Fc Fab

Some activated B cells do not differentiate into plasma cells but instead become memory B cells (discussed in a later section).

A typical antibody consists of four polypeptide chains An antibody molecule, also called immunoglobulin (Ig), has several functions. It combines with antigen, and it activates processes that destroy the antigen that binds to it. For example, an antibody may stimulate phagocytosis. Note that an antibody does not destroy an antigen directly; rather, it labels the antigen for destruction. The antibody molecule also serves as the B-cell receptor for antigen when it is attached to the B-cell surface. Rodney Porter, of the University of Oxford in England, and Gerald Edelman, of Rockefeller University in New York, clarified the basic structure of the immunoglobulin molecule during the 1960s. Porter used the plant enzyme papain, a protease, to split Ig molecules into fragments. Based on his findings, Porter developed a working model of the structure of the Ig molecule and was the first to suggest that it is Y shaped. These researchers then constructed an accurate model of the antibody molecule. Porter and Edelman were awarded the 1972 Nobel Prize in Medicine for their contributions. In a Y-shaped antibody molecule, the tail of the Y is called the Fc fragment (c indicates that this fragment crystallizes during cold storage). The Fc fragment binds with receptors on phagocytes and other cells of the immune system. This region of the antibody molecule also binds with molecules of the complement system. The two arms of the Y are referred to as Fab fragments (Fab stands for fragment, antigen-binding). The tips of the Fab fragments bind with antigen (FIG. 45-9). The Y shape permits the antibody molecule to combine with two antigen molecules, which allows the formation of antigen–antibody complexes. Each antibody molecule consists of four polypeptide chains: two long chains, called heavy chains, and two short chains, called light chains. Each chain has a constant region and a variable region. In the constant (C) region of the heavy chains, the amino acid sequence is constant within a particular immunoglobulin class. The C region of the heavy chains is the tail of the Y part of the antibody molecule. You can think of this C region as the handle portion of a door key. The variable (V) region has a unique amino acid sequence, like the pattern of bumps and notches on the part of a key that fits into a lock. At its variable regions, the antibody folds three-

Antigen–antibody complex Inactivates pathogen

Inactivates toxins

Increases phagocytosis Activates complement

Destroys pathogen

Coats surface of pathogen

Increases phagocytosis

Stimulates mast cells to release histamine

Attracts phagocytes

Increases inflammation

(b) The Fab portion of the antibody binds to antigen. The Fc portion of the molecule binds to a cell of the immune system (not shown). Antigen–antibody complexes directly inactivate pathogens and increase phagocytosis. They also activate the complement system.

FIGURE 45-9 Animated Antibody structure and function Antibodies combine with antigens to form antigen–antibody complexes. These complexes destroy or promote the destruction of pathogens in a variety of ways.

dimensionally, assuming a shape that enables it to combine with a specific antigen. When they meet, antigen and antibody fit together somewhat like a lock and key. They must fit in just the right way for the antibody to be effective (FIG. 45-10).

Both: Fig. 3AB from Amit et al., Science 233: 747–753 (1986)

(a) A portion of the antigenic determinant (red ) fits into a groove in the antibody molecule.

(b) The antigen–antibody complex has been pulled apart to show its structure.

FIGURE 45-10 An antigen–antibody complex The components of an antigen–antibody complex fit together as shown in this computer simulation of their molecular structure. Antigen lysozyme, green; heavy chain of the antibody, blue; light chain, yellow.

The specific part of the antigen molecule that is recognized by an antibody or T-cell receptor is called an antigenic determinant, or epitope. An antigen may have many antigenic determinants on its surface; some have hundreds. Antigenic determinants often differ from one another, so several different kinds of antibodies can combine with a single antigen. A given antibody can bind with different strengths, or affinities, to different antigens. In the course of an immune response, higher affinity antibodies are generated. This occurs because as activated B cells are replicated, mutations occur in genes that encode the antibodies. Cells with antibodies that bind more tightly are selected during the ongoing immune response—a striking example of microevolution in action.

signaling molecules. These chemicals are released when an antigen binds to IgE on a mast cell. Histamine triggers many allergy symptoms, including inflammation. IgE is also responsible for an immune response to invading parasitic worms.

Antibodies are grouped in five classes

1. The antigen–antibody complex may inactivate the pathogen or its toxin. For example, when an antibody attaches to its surface, a virus may lose the ability to attach to a host cell. 2. The antigen–antibody complex stimulates phagocytic cells to ingest the pathogen. 3. Antibodies of the IgG and IgM groups work mainly through the complement system.

Antibodies are grouped in five classes, defined by unique amino acid sequences in the C regions of the heavy chains. The classes are named using the abbreviation Ig for immunoglobulin and are designated IgG, IgM, IgA, IgD, and IgE. In humans, about 75% of the circulating antibodies belong to the IgG class; these are part of the gamma globulin fraction of the plasma. IgM is the first antibody that humans make in an immune response. IgG and IgM share some functional properties. Both interact with macrophages and activate the complement system. They defend against many pathogens carried in the blood, including bacteria, viruses, and some fungi. IgG is the antibody that crosses the placenta (the organ of exchange between mother and developing fetus) and protects the fetus and, later, the newborn baby. IgA, present in mucus, tears, saliva, and breast milk, prevents viruses and bacteria from attaching to epithelial surfaces. This immunoglobulin, which defends against inhaled or ingested pathogens, is strategically secreted into the respiratory, digestive, urinary, and reproductive tracts. Like IgM, IgD is an important immunoglobulin on the B-cell surface. It has a critical role in maturation of B cells and helps activate B cells following antigen binding. IgD has a low concentration in plasma (less than 1% of circulating antibodies). IgE, which has an even lower plasma concentration than IgD, binds to mast cells, which contain histamine and other potent

Antigen–antibody binding activates other defenses Antibodies mark a pathogen as foreign by combining with an antigen on its surface. Generally, several antibodies bind with several antigens, creating a mass of clumped antigen–antibody complexes. The combination of antigen and antibody activates several defensive responses (see Fig. 45-9).

When antibodies combine with a specific antigen on a pathogen, complement proteins destroy the pathogen. IgG molecules have an Fc fragment that binds Fc receptors; these receptors are expressed on many cells, including macrophages and neutrophils. When an antibody molecule is bound to a pathogen and the Fc portion of the antibody binds with an Fc receptor on a phagocyte, the pathogen is more easily destroyed.

The immune system responds to millions of different antigens How can the immune system recognize every possible antigen, even those produced by newly mutated viruses never before encountered during the evolution of our species? In the 1950s, the Danish researcher Niels Jerne, of the Basel Institute for Immunology in Basel, Switzerland; David Talmage, of the University of Chicago; and Frank Macfarlane Burnet, of the Walter and Eliza Hall

Institute for Medical Research in Melbourne, Australia, developed the clonal selection hypothesis. This hypothesis proposes that before a lymphocyte ever encounters an antigen, the lymphocyte has specific receptors for that antigen on its surface. ( Jerne and Burnet later were awarded Nobel Prizes in Physiology or Medicine for their work in immunology.) When an antigen binds to a matching receptor on a lymphocyte, it activates the lymphocyte, which then gives rise to a clone of cells with identical receptors. A problem with this hypothesis when it was initially proposed was its suggestion that our cells must contain millions of separate antibody genes, each coding for an antibody with a different specificity. Although each human cell has a large amount of DNA, it is not enough to provide a different gene to encode each of the millions of possible specific antibody molecules. An explanation for how the immune system could have so many different specificities came later. In 1965, W. J. Dreyer of the Institute of Biochemistry at the University of Zurich, Switzerland, and Joe Claude Bennett of the University of Alabama at Birmingham suggested that the C region and the V region of an immunoglobulin are encoded by two separate genes. Their hypothesis was at first rejected by many biologists because it contradicted the prevailing hypothesis that one gene codes for one polypeptide. The technology needed to test Dreyer and Bennett’s hypothesis was not immediately available, and it was not until 1976 that Susumu Tonegawa of the Massachusetts Institute of Technology and his colleagues demonstrated that separate genes encode the V and C regions of immunoglobulins. Tonegawa further showed that three separate families of genes code for immunoglobulins and that each gene family contains a large number of DNA segments that code for V regions. Recombination of these DNA segments during the differentiation of B cells is responsible for antibody diversity. Tonegawa was awarded the 1987 Nobel Prize in Physiology or Medicine for his work, which transformed the emerging science of immunology. We now understand that in undifferentiated B cells, gene segments are present for a number of different V regions, for one or more junction ( J) regions, and for one or more different C regions (FIG. 45-11). Rearrangement of these DNA segments produces an enormous number of potential combinations. Millions of different types of B (and T) cells are produced. By chance, one of those B cells may produce just the right antibody to destroy a pathogen that invades the body. To appreciate the capacity for antibody diversity, consider the diverse combinations of things you create in your everyday life. A familiar example is making an ice cream sundae. Imagine the varied combinations that are possible using 10 flavors of ice cream, 6 types of sauce, and 15 kinds of toppings. Researchers have identified additional sources of antibody diversity. For example, the DNA of the mature B cells that codes for the regions of immunoglobulins mutates very readily. These somatic mutations produce genes that code for slightly different antibodies. We have used antibody diversity here as an example, but similar genetic mechanisms account for the diversity of T-cell receptors. Although we may actually use only a relatively few types of antibodies or T cells in a lifetime, the remarkable diversity of the immune system prepares it to attack most potentially harmful antigens that may invade the body.

Monoclonal antibodies are highly specific Before 1975, the only method for obtaining antibodies for medicine or research was immunizing animals and collecting their blood. Then Cesar Milstein and Georges Kohler at the Laboratory of Molecular Biology in Cambridge, England, developed monoclonal antibodies, identical antibodies produced by cells cloned from a single cell. One way to produce monoclonal antibodies in the laboratory is to inject mice with the antigen of interest, for example, an antigen from a particular bacterium. After the mice have produced antibodies to the antigen, their plasma cells are collected. However, these normal plasma cells survive in culture for only a few generations, limiting the amount of antibody that can be produced. In contrast, cancer cells live and divide in tissue culture indefinitely. B cells can be suspended in a culture medium together with lymphoma cells from other mice. (Lymphoma is a cancer of lymphocytes.) Then the B cells and lymphoma cells are induced to fuse. They form hybrid cells, known as hybridomas, which have properties of the two “parent” cells. Hybridomas can be cultured indefinitely (a cancer cell property) and continue to secrete antibodies (a B-cell property). Researchers select hybrid cells that are manufacturing the specific antibody needed and then clone them in a separate cell culture. Cells of this clone secrete large amounts of the particular antibody—thus the name monoclonal antibodies. Each type of monoclonal antibody is specific for a single antigenic determinant. Immunologists also use recombinant DNA technology to produce monoclonal antibodies. Because of their purity and specificity, monoclonal antibodies have proved to be invaluable tools in modern biology. For example, a researcher may want to detect a specific molecule present in very small amounts in a mixture. The reaction of that molecule (the antigen) with a specific monoclonal antibody makes its presence known. Monoclonal antibodies are used in similar ways in various diagnostic tests. Home pregnancy tests make use of a monoclonal antibody that is specific for human chorionic gonadotropin (hCG), a hormone produced by a developing human embryo (see Chapters 50 and 51). Several monoclonal antibodies are being used clinically to treat cancer. The drug Herceptin is a monoclonal antibody used in treating a form of breast cancer.

Immunological memory is responsible for long-term immunity Immunological memory depends on memory B cells and memory T cells. Following an immune response, memory T cells group strategically in many nonlymphatic tissues, including the lung, liver, kidney, and intestine. Many infections occur at such sites, and T cells stationed in those locations can respond quickly. In response to antigen, memory T cells rapidly become TC cells and produce substances that kill invading cells. Memory B cells have a “survival gene” that prevents apoptosis, the programmed cell death that is the eventual fate of plasma cells. Memory B cells continue to live and produce small amounts of antibody long after the body has overcome an infection. This circulating antibody is part of the body’s arsenal of chemical weapons.

DNA of stem cell V1

V2

V3

V4

J1

J3

J2

Intron

C

Gene rearrangement V1

V2

V3 J3

Intron

C

B-cell DNA Functional gene Transcription V3 RNA transcript

J3

Intron

C

5′

3′

RNA splicing V3 J3

C

5′

3′ Translation

Polypeptide Variable region

Constant region Antibody molecule

FIGURE 45-11 Animated Antibody diversity In undifferentiated cells, gene segments are present for several different variable (V) regions, for one or more junction (J) regions, and for one or more different constant (C) regions. During differentiation, these DNA segments become rearranged. A gene segment extending from the end of one V segment to the beginning of one J segment may be deleted, thus producing a gene that can be transcribed. The RNA transcript is processed to remove introns, and the mRNA is translated. Each of the polypeptide chains of an antibody molecule is produced in this way.

If the same pathogen enters the body again, the antibody immediately targets it for destruction. At the same time, specific memory cells are stimulated to divide, producing new clones of plasma cells that produce the same antibody. The presence of circulating antibodies is used clinically to detect previous exposure to specific pathogens. For example, some HIV screening tests measure antibody to the virus that causes AIDS. A secondary immune response is more effective than a primary response The first exposure to an antigen stimulates a primary immune response. An infection, or an antigen injected into an animal, causes specific antibodies to appear in the blood plasma in 3 to 14 days. After antigen injection, there is a brief latent period during which

the antigen is recognized and appropriate lymphocytes begin to form clones. A logarithmic phase follows, during which the antibody concentration rises rapidly for several days until it reaches a peak (FIG. 45-12). IgM is the principal antibody synthesized during the primary immune response. Finally, there is a decline phase, during which the antibody concentration decreases to a very low level. A second exposure to the same antigen, even years later, results in a secondary immune response. Because memory B cells bearing antibodies to that antigen (and also memory T cells with their specific T-cell receptors) persist for many years, the secondary immune response is much more rapid than the primary immune response and has a shorter latent period. Much less antigen is necessary to stimulate a secondary immune response than a primary response, and more antibodies are produced. In addition, the

Antibodies to antigen 1—primary response Antibodies to antigen 1—secondary response Antibodies to antigen 2—primary response

Antibody concentration (arbitrary units)

104

change may prevent recognition by memory cells. Because the immune system is so specific, each different antigen is treated by the body as a new immunological challenge. Immunization induces active immunity

103 102

Inject antigens 1 and 2

Inject antigen 1

101

0 0

1

2

3

4

5

6

7

8

Time (weeks)

FIGURE 45-12 Animated Immunological memory Antigen 1 was injected at day 0, and the immune response was assessed by measuring levels of antibody specific for antigen 1. At week 4, the primary immune response had subsided. Antigen 1 was injected again, together with a new protein, antigen 2. The secondary immune response was greater and more rapid than the primary response. It was also specific to antigen 1. A primary immune response was made to the newly encountered antigen 2.

affinity of antibodies is generally much higher. The predominant antibody in a secondary immune response is IgG. The body’s ability to launch a rapid, effective response during a second encounter with an antigen explains why we do not usually suffer from the same infectious disease several times. For example, a person who contracts measles or chickenpox is unlikely to become re-infected in the future. When someone is exposed a second time, the immune system responds quickly, destroying the pathogens before they have time to multiply and cause symptoms of the disease. Booster shots of vaccine are given to elicit a secondary immune response, thus reinforcing immunological memory. You may wonder how a person can get influenza (the flu) or a cold more than once. The reason is that there are many varieties of these viral diseases, each caused by a virus with slightly different antigens. For example, more than 100 different viruses cause the common cold, and new varieties of cold and flu viruses evolve continuously by mutation (a survival mechanism for them). As viruses evolve, changes may occur in their surface antigens. Even a slight TABLE 45-1

We have been considering active immunity, which develops following exposure to antigens. Active immunity can be naturally or artificially induced (TABLE 45-1). If someone with measles sneezes near you and you contract the disease, you develop active immunity naturally. Active immunity can also be artificially induced by immunization, that is, by exposure to antigens in a vaccine. When an effective vaccine is introduced into the body, the immune system actively develops clones of cells, produces antibodies, and develops memory cells. The first vaccine was prepared in 1796 by British physician Edward Jenner against vaccinia, the cowpox virus. (The term vaccination was derived from the name of the cowpox virus.) Jenner’s vaccine provided humans with immunity against the deadly disease smallpox, which closely resembles cowpox. Jenner had no knowledge of microorganisms or of immunology, and 100 years passed before French chemist Louis Pasteur began to develop scientific methods for preparing vaccines. Pasteur showed that inoculations with preparations of attenuated (weakened) pathogens could be used to develop immunity against the virulent (infectious) form of the pathogen. However, not until 20th-century advances in immunology—for example, Burnet’s clonal selection theory in 1957 and the discovery of T cells and B cells in 1965—did scientists gain a modern understanding of vaccines. Effective vaccination stimulates the body to launch an immune response against the antigens contained in the vaccine. Memory cells develop, and future encounters with the same pathogen are dealt with rapidly. Microbiologists prepare effective vaccines in various ways. They can attenuate a pathogen so it loses its ability to cause disease. When pathogens are cultured for long periods in nonhuman cells, mutations adapt the pathogens to the nonhuman host so that they no longer cause disease in humans. This method is used to produce the Sabin polio, measles, mumps, and rubella vaccines. Tetanus and diphtheria vaccines are made from toxins secreted by the respective pathogens. The toxin is altered so that it can no longer destroy tissues, but its antigenic determinants are still intact. Researchers are developing DNA vaccines (or RNA vaccines) made from a part of the pathogen’s genetic material. The DNA of

Active and Passive Immunity Development of Memory Cells

Duration of Immunity

Yes

Many years

Artificially induced

After pathogens enter the body through natural encounters (e.g., a person with measles sneezes on you) After immunization with a vaccine

Yes

Many years

Passive Naturally induced Artificially induced

After transfer of antibodies from mother to developing baby After injection with gamma globulin

No No

Few months Few months

Type of Immunity Active Naturally induced

When Developed

the pathogen is altered so that it transfers genes that specify antigens. When injected into a patient, the altered DNA is taken up by cells and makes its way to the nucleus. The encoded antigens are manufactured and stimulate both cell-mediated and antibodymediated immunity. Several DNA vaccines, including vaccines to prevent and treat HIV infection, are in clinical trials. Passive immunity is borrowed immunity To provide passive immunity, physicians inject people with antibodies actively produced by another organism. The serum or gamma globulin that contains these antibodies is obtained from humans or other animals. Nonhuman serums are less desirable because nonhuman proteins can act as antigens, stimulating an immune response that may result in serum sickness. Passive immunity is borrowed immunity. Its effects are temporary whether the immunity is artificially or naturally induced (see Table 45-1). Physicians use artificially induced passive immunity to boost the body’s defenses temporarily against a particular disease. For example, when exposed to hepatitis A, a form of viral hepatitis spread through contaminated food or water, people at risk of infection may be injected with gamma globulin containing antibodies to the hepatitis pathogen. However, gamma globulin injections offer protection for only a few weeks. Because the body has not actively launched an immune response, it has no memory cells and cannot produce antibodies to the pathogen. Once the injected antibodies are broken down, the immunity disappears. Pregnant women confer naturally induced passive immunity on their developing babies by manufacturing antibodies for them. These maternal antibodies, of the IgG class, pass through the placenta and provide the fetus and newborn infant with a defense system until its own immune system matures. Babies who are breast-fed continue to receive immunoglobulins, particularly IgA, in their milk, during the critical months when their own immune system is developing.

Review ■ ■

■ ■

How do antibodies recognize antigens? What are three ways that antigen–antibody complexes affect pathogens? What is the basis of immunological memory? How is passive immunity different from active immunity?

45.6 RESPONSE TO DISEASE, IMMUNE FAILURES, AND HARMFUL REACTIONS ■ ■ LEARNING OBJECTIVES 11 Describe the body’s response to cancer cells and HIV. 12 Describe examples of hypersensitivity, including autoimmune diseases, Rh incompatibility, and allergic reactions, and summarize the immunological basis of graft rejection.

The immune system is generally very effective in defending the body against invading pathogens. However, many factors can im-

pair immune function, including genetic mutations, malnutrition, sleep deprivation, pre-existing disease, stress, and virulent pathogens. Ineffective or inappropriately directed immune responses can also result in disease states. Exaggerated, damaging immune responses are referred to as hypersensitivity. For example, sometimes the immune system malfunctions, and attacks its own tissues, causing autoimmune disease. In allergic reactions, individuals are overresponsive to mild environmental antigens. The immune system’s ability to distinguish self from nonself sometimes interferes with clinical interventions. Physicians must carefully consider immune system responses when employing such life-saving procedures as blood transfusions or organ transplants.

Cancer cells evade the immune system Cancer is a leading cause of death worldwide and the secondleading cause of death in Western countries. An estimated one person in three in the United States will develop cancer, and one in five will die of cancer. Worldwide, lung cancer in men and breast cancer in women are the main types of cancer resulting in death. (In our earlier discussions of cancer in Chapters 17 and 39, we explained that cancer is actually a complex group of diseases characterized by dysregulation of normal control mechanisms for cellular growth.) Here we focus on the interaction between cancer cells and cells of the immune system and on new treatment strategies based on immune mechanisms. Cancer cells are body cells that have been transformed in a way that alters their normal growth-regulating mechanisms. Every day, a few normal cells may be transformed into precancer cells in each of us in response to the sun’s UV rays, X-rays, certain viruses, tobacco smoke, asbestos, and other chemical carcinogens in the environment, as well as other unknown factors. How does the immune system respond? And why is immune response sometimes ineffective? Cancer cell antigens are classified according to their molecular structure and their source. Some cancer cell antigens are produced by mutant genes that are associated with cancer cells. T cells may respond to them because these antigens are not found in normal cells. Other cancer cell antigens are normal proteins that may stimulate immune responses because they are expressed in abnormally large quantities. The human papillomavirus (HPV), which has been linked to cervical cancer, produces antigens that stimulate T cells to respond. Specific immune responses sometimes prevent the development of such DNA virus-induced cancers. Cancer cell antigens induce both cell-mediated and antibody-mediated immunity NK cells and macrophages destroy cancer cells. Macrophages produce cytokines, including TNFs that inhibit tumor growth. Dendritic cells present antigens to T cells, stimulating them to produce interferons, which have an antitumor effect. TC cells attack cancer cells directly and also produce interleukins, which attract and activate macrophages and NK cells (FIG. 45-13). Although the immune system recognizes many cancer cell antigens, these antigens often stimulate only weak immune responses that are not effective.

(a) An army of TC cells surrounds a large cancer cell. The TC cells recognize the cancer cell as nonself because it displays altered or unique antigens on its surface.

Lennart Nilsson, Boehringer Ingelheim International GmbH

Lennart Nilsson, Boehringer Ingelheim International GmbH

Lennart Nilsson, Boehringer Ingelheim International GmbH

10 μm

10 μm

10 μm

(b) Some TC cells elongate as they chemically attack the cancer cell and break down its plasma membrane.

(c) The cancer cell has been destroyed. Only a collapsed fibrous cytoskeleton remains.

FIGURE 45-13 Destroying cancer cells This series of colorized SEMs shows TC cells (green) attacking and destroying a cancer cell (yellow).

Sometimes cancer cells evade the immune system and multiply unchecked. Some types of cancer cells can block TC cells. Others dramatically decrease their expression of class I MHC molecules. Recall that TC cells recognize only antigens associated with class I MHC, so a low level of these molecules prevents tumor destruction. Some cancer cells do not produce co-stimulatory molecules needed to activate TC cells. Future cancer treatment will be more targeted Conventional cancer treatments, such as chemotherapy and radiation, are designed to kill cells that are rapidly dividing. However, these treatments destroy normal cells as well as cancer cells. To treat cancer effectively, treatment strategies must be more specific, or even customized to the particular cancer. For example, cancer researchers are genetically engineering cancer cells to secrete cytokines that would stimulate the patient’s immune response. A few cancer drugs, such as Herceptin, are monoclonal antibodies. Herceptin binds with a growth-factor receptor that is present in excessive numbers on the cells of about 30% of metastatic breast cancers. Herceptin blocks the growth factors that would stimulate proliferation of the cells. The spread of cancer typically depends on the development of blood vessels that bring nutrients and oxygen to the tumor. Angiogenesis inhibitors slow the development of blood vessels, and thereby slow the spread of cancer. Many studies underway are attempting to optimize the clinical benefits of these drugs. Researchers are developing cancer vaccines that contain cancer cells or tumor antigens; they stimulate the patient’s immune system to launch cell-mediated responses to tumor-associated antigens. These vaccines differ from traditional vaccines that are

used to destroy pathogens before they cause disease. DNA vaccines are made from bits of DNA from a patient’s cancer cells. When injected into the patient, the vaccine stimulates increased antigen production. In response, the patient’s immune system produces more T cells. Investigators are making dendritic cell vaccines by taking dendritic cells from a cancer patient and exposing them to antigens from the patient’s cancer cells. The combination of dendritic cells and antigens is then injected into the patient. The dendritic cells display the antigens to TC cells, which then attack the cancer cells. Another approach that has recently shown promise is the development of drugs that deactivate proteins that activate key signaling pathways. Effective treatment protocols for cancer will almost certainly utilize a systems approach and combinations of medications. Cellular mechanisms are complex, and when a drug blocks one signaling pathway, cancer cells can work around the blocked reaction or take advantage of a redundant pathway. When combinations of drugs are used, more than one signaling pathway, or different parts of the same pathway, can be targeted.

Immunodeficiency disease can be inherited or acquired The absence or failure of some component of the immune system can result in immunodeficiency disease, a condition that increases susceptibility to infection. Inherited immunodeficiencies have an estimated incidence of 1 per 10,000 births. Severe combined immunodeficiency syndromes (SCIDs) are X-linked and autosomal recessive disorders that profoundly affect both cell-mediated immunity

and antibody-mediated immunity, resulting in multiple infections. Babies born with SCID typically die by two years of age unless they are maintained in a protective bubble until they can be effectively treated, typically with bone marrow transplants. In DiGeorge syndrome, the thymus is reduced or absent and the patient is deficient in T cells. Children born with this disorder are prone to serious viral infections. Treatment involves transplanting bone marrow or fetal thymus tissue. Other tissues may take over Tcell production, and function typically improves by age five years. Researchers use several animal models to study immunodeficiency. For example, the nude mouse (a hairless mutant mouse) does not develop a functional thymus. Because nude mice are deficient in mature T cells, they are used to study the effects of compromised cell-mediated immunity, as well as possible treatments. Stem cell research and advances in genetic engineering suggest new approaches to treating immunodeficiency (see Chapters 16 and 17). Worldwide, the leading cause of acquired immunodeficiency in children is protein malnutrition. Lack of protein decreases Tcell numbers and decreases the ability to manufacture antibodies, which impairs the immune system. This condition increases the risk of contracting opportunistic infections, infections by microbes that do not typically cause disease. Another important cause of acquired immunodeficiency is chemotherapy administered to cancer patients because chemotherapy is most damaging to cells that divide rapidly, including blood stem cells and activated lymphocytes.

HIV is the major cause of acquired immunodeficiency in adults The human immunodeficiency virus (HIV) was first isolated in 1983 and was shown to be the cause of acquired immunodeficiency syndrome (AIDS) in 1984. HIV, which is a retrovirus, has been studied more than any other virus. (Recall from Chapter 24 that a retrovirus is an RNA virus that uses its RNA as a template to make DNA with the help of reverse transcriptase.) Several different strains of the virus are known; HIV-1 is the most virulent form in humans. The AIDS pandemic has claimed more than 30 million lives. HIV kills more than 2 million people every year, making it the sixth-highest cause of death globally. Epidemiologists estimate that more than 33 million people, including more than 2 million children, worldwide are now infected with HIV. More than 2.7 million people are newly infected with HIV each year. The General Assembly of the United Nations has called AIDS a “global emergency” and has outlined measures for mobilizing world resources to combat this disease. HIV/AIDS is more than a medical issue; it is a political, economic, and human rights threat. HIV is transmitted mainly during sexual intercourse with an infected person or by direct exposure to infected blood or blood products. Sharing needles and/or syringes (usually for drug injection) with an infected person is a major risk factor. The virus is not spread by casual contact. People do not contract HIV by hugging, casual kissing, or using the same bathroom facilities. Friends and family members who have contact with AIDS patients are not more likely to become infected. In the United States, more than 50,000 new HIV infections are occurring each year. Of these new infections, about 75% occur in

men. Sexual contact with persons who have HIV infection is a risk factor that can be modified. Use of a latex condom during sexual intercourse provides some protection against HIV transmission, and use of a spermicide containing nonoxynol-9 may provide additional protection. As many as one-third of untreated pregnant women infected with HIV will transmit the virus to their babies during pregnancy, childbirth, or breast-feeding. Recent drug treatment protocols have reduced the rate of HIV transmission between mother and fetus. In highly developed countries, effective blood-screening procedures now safeguard blood bank supplies, markedly reducing the risk of infection by blood transfusion. HIV destroys the host immune system HIV typically enters the body through the mucosa that lines passageways. A protein on the outer envelope of the virus attaches to CD4 on the surface of TH cells, which are the main target (FIG. 45-14; also see Chapter 23 opening photograph). The virus also binds to a coreceptor that normally binds a chemokine. HIV enters the TH cell and is reverse-transcribed to form DNA. Its DNA is then incorporated into the host genome. The virus reproduces itself, causing death of the host cell. The first phase of HIV infection is the acute phase of the infection. Within two weeks of infection, large numbers of TH and memory T cells are destroyed. HIV also infects dendritic cells and macrophages. Dendritic cells in the mucous membranes transport HIV from their point of entry to the lymph nodes. The virus further spreads through the blood. The body launches an immune response following HIV infection. Nonspecific responses include production of antimicrobial peptides and action by NK cells, dendritic cells, and the complement system. Specific immune responses include activation and mobilization of TC cells. An estimated 25% of HIV-infected individuals in the United States do not know they are infected. Many individuals experience mild flulike symptoms (fever, headache, sore throat, swollen lymph glands, aching muscles) about two weeks after contracting the virus. However, some individuals have no symptoms. The acute phase transitions into the latent period, a chronic phase, as the immune system decreases HIV in the blood. Some cells infected by HIV are not destroyed, and the virus may continue to replicate slowly for many years. TC cells appear to be the main cells that attack HIV, limiting viral replication and delaying the progress of the disease. However, over time HIV continues to destroy TH cells, resulting in a dramatic decrease in the TH cell population. This decrease severely impairs the body’s ability to resist infection because there are no longer TH cells to release the cytokines necessary to activate TC cells and B cells. After a time, a progression of symptoms occurs, including swollen lymph glands, night sweats, fever, and weight loss (FIG. 45-15). Too often, the virus eventually wins the battle. In most HIV-infected individuals, the disease eventually progresses to AIDS, the final, and often lethal, stage. This process can take more than ten years, and antiretroviral drugs slow the process. By the time the disease progresses to AIDS, lymph tissue has been destroyed and the blood TH cell count has dropped below a critical level of 200 cells/mL. As a consequence of this immunosuppression, many AIDS patients develop serious opportunis-

Exposure to semen or blood containing HIV Certain sexually transmitted diseases

Level of immune function

Psychosocial factors

Lennart Nilsson, Boehringer Ingelheim International GmbH

Genetic factors

No infection

Other environmental factors

HIV infection; acute phase

Latent period

Progression of symptoms

300 nm

(a) HIV (blue particles) attacking TH cells (light green).

Swollen lymph glands, night sweats, fever, weight loss

AIDS

Severely depressed immune function

AIDS dementia complex

Lennart Nilsson, Boehringer Ingelheim International GmbH

Opportunistic infections

Death

Death

FIGURE 45-15 Animated The course of HIV infection

100 nm

(b) HIV (purple/blue) budding from the ends of the branched microvilli of a TH cell.

Exposure to semen or blood that contains HIV can lead to HIV infection. The acute (early) infection begins in lymph nodes and other lymph tissues associated with epithelial linings. Most TH cells may be destroyed within two weeks of infection. During the latent period, chronic infection develops and the virus replicates at a low level. Eventually, viral replication may increase and lead to a progression of symptoms and to AIDS, the final, often lethal, stage. Although some exposed individuals apparently do not become infected, the risk increases with multiple exposures. Many factors determine whether a person exposed to HIV will develop AIDS.

HIV infection depletes TH lymphocytes. This is the main reason for the failure of the immune system in AIDS.

AIDS dementia complex. These patients exhibit progressive cognitive, motor, and behavioral dysfunction that typically ends in coma and death.

tic infections, such as tuberculosis, or rare forms of cancer, such as Kaposi’s sarcoma (a tumor of the endothelial cells lining blood vessels that causes purplish spots on the skin). In about one-third of AIDS patients, the virus infects the nervous system and causes

Researchers are searching for effective treatments Researchers throughout the world are searching for drugs that will successfully combat the AIDS virus. Because HIV often infects the central nervous system, an effective drug must cross the blood– brain barrier. AZT (azidothymidine), the first drug developed to

FIGURE 45-14 Colorized SEMS of HIV infecting T helper cells

treat HIV infection, can prolong the period prior to the onset of AIDS symptoms. AZT inhibits the action of reverse transcriptase, the enzyme the retrovirus uses to synthesize DNA. As a result, AZT blocks HIV replication. Without producing DNA, the virus cannot incorporate itself into the host cell’s DNA. Unfortunately, the reverse transcriptase HIV uses to synthesize DNA makes many mistakes: about 1 in every 2000 nucleotides it incorporates is incorrect. By mutating in this way, viral strains resistant to AZT have evolved. Protease inhibitors block the viral enzyme protease, resulting in viral copies that cannot infect new cells. Protease inhibitors are used in combination with AZT and other reverse transcriptase inhibitors. Triple-combination treatment has been very effective for many AIDS patients, preventing opportunistic infections and prolonging life. In fact, combination treatment has led to a decline in AIDS incidence and mortality in the United States. Unfortunately, people in many parts of the world cannot afford these expensive drugs, and in impoverished areas, the AIDS epidemic continues to grow. In addition, the emergence of drug-resistant HIV viruses has been a serious problem. Worldwide containment of the AIDS epidemic will require an effective vaccine that prevents the spread of HIV. Developing such a vaccine remains a daunting challenge for immunologists. More than 50 vaccines have been developed and tested clinically, but none has proved effective. An effective vaccine would have to counteract HIV’s destruction of key cells needed to mount an immune response. Also, because mutations arise at a very high rate, new viral strains with new antigens evolve quickly. A vaccine would not be effective against new antigens and would quickly become obsolete. Some optimism has been generated by an apparently effective combination of vaccine and gene therapy that has been tested in simian monkeys. Also, recent human clinical trials that combined two vaccines yielded marginally positive results. While immunologists work to develop a successful vaccine and more effective drugs to treat infected patients, other strategies are being utilized with some success. For example, we now know that male circumcision reduces the risk of HIV infection through sexual contact. Massive educational programs have been launched to slow the spread of HIV. Educating the public that having multiple sexual partners increases the risk of AIDS and teaching sexually active individuals the importance of “safe sex” may help contain the pandemic. Some have suggested that public health facilities should provide free condoms to those who are sexually active and free sterile hypodermic needles to those addicted to intravenous drugs. The cost of these measures would be far less than the cost of medical care for increasing numbers of AIDS patients and the toll in human suffering.

In an autoimmune disease, the body attacks its own tissues Self-tolerance, the ability to recognize self, is established during lymphocyte development. Lymphocytes do not normally attack cells and tissues of their own body. However, some lymphocytes have the potential to be autoreactive, that is, to launch an immune response against self-tissues. In about 5% of adults in highly devel-

oped countries, such autoreactivity can lead to a form of hypersensitivity known as autoimmunity, or autoimmune disease, in which immune cells react against the body’s own cells. Some of the diseases that result from such failure in self-tolerance are rheumatoid arthritis, systemic lupus erythematosus (SLE), insulin-dependent diabetes, scleroderma, and multiple sclerosis. In autoimmune diseases, antibodies and T cells attack the body’s own tissues. In rheumatoid arthritis, T cells in the area of inflammation produce a cytokine that promotes inflammation. In addition, antibodies to a joint protein may generate antigen– antibody complexes that are deposited in the small joints. In multiple sclerosis, an autoimmune disease of the central nervous system, TH cells attack self-myelin antigens. MRI scans show the loss of myelin sheaths in multiple areas around axons in the brain and spinal cord that are normally myelinated. Scar tissue forms and compromises the ability of these damaged neurons to transmit signals. Multiple genes, including MHC genes, contribute to the development of autoimmune disease. Mutations that cause abnormal apoptosis or inadequate clearing of dead cells may result in danger signals that activate the immune system. Another important factor is that viral or bacterial infection often precedes the onset of an autoimmune disease. Some pathogens have evolved a tactic known as molecular mimicry. They trick the body by producing molecules that look like self-molecules. For example, an adenovirus that causes respiratory and intestinal illness produces a peptide that mimics myelin protein. When the body launches responses to the adenovirus peptide, it may also begin to attack the similar self-molecule, myelin. To maintain homeostasis and health, the numerous types of cells, regulatory and signaling molecules, and the complex actions of the immune system must be carefully regulated. Specific T cells and B cells proliferate and respond to challenges by pathogens, but after an effective immune response the expanded populations of T cells and B cells must be decreased to normal numbers. Cells that are no longer needed die by apoptosis. Overactivation of critical components can result in inflammation and autoimmune disease. Underactivation allows pathogens to cause serious disease. Either situation can lead to death.

Rh incompatibility can result in hypersensitivity Named for the rhesus monkeys in whose blood it was first found, the Rh system consists of more than 40 kinds of Rh antigens, each referred to as an Rh factor. By far the most important of these factors is antigen D. About 85% of U.S. residents who are of Western European descent are Rh positive—they have antigen D on the surfaces of their red blood cells (in addition to the antigens of the ABO system and other blood group antigens). The 15% or so of this population who are Rh negative have no antigen D. Unlike the situation discussed in Chapter 11 for the ABO blood group, Rh-negative individuals do not naturally produce antibodies against antigen D (anti-D). However, they do produce anti-D antibodies if they are exposed to Rh-positive blood. The allele coding for antigen D is dominant to the allele for the absence of antigen D. For this reason, Rh-negative people are homozygous recessive, and Rh-positive people are heterozygous or homozygous dominant.

Although several kinds of maternal-fetal blood type incompatibilities are known, Rh incompatibility is the most serious (FIG. 45-16). If a woman is Rh negative and the father of the fetus she is carrying is Rh positive, the fetus may also be Rh positive, having inherited the D allele from the father. Ordinarily, the maternal and fetal blood do not mix; nutrients, oxygen, and other substances are exchanged between these two circulatory systems across the placenta. However, late in pregnancy or during the birth process, a small quantity of blood from the fetus may pass through a defect in the placenta. If any maternal blood mixes with fetal blood, fetal red blood cells, which bear antigen D, activate the mother’s immune system. The mother’s B cells then produce antibodies to antigen D. If the woman becomes pregnant again, anti-D antibodies cross the placenta and enter the fetal blood. They combine with antigen D molecules on the surface of the fetal red blood cells, causing hemolysis (cells rupture and release hemoglobin into the circulation). The ability to transport oxygen is reduced, and breakdown products of the released hemoglobin damage organs, including the brain. In extreme cases of this disease, known as erythroblastosis fetalis, so many fetal red blood cells are destroyed that the fetus may die. When Rh-incompatibility problems are suspected, fetal blood can be exchanged by transfusion before birth, but this is a risky procedure. More commonly, Rh-negative women are treated just after childbirth (or at termination of pregnancy by miscarriage

or abortion) with a preparation of anti-D antibodies known as RhoGAM. These antibodies clear the Rh-positive fetal red blood cells from the mother’s blood very quickly, minimizing the chance for her own white blood cells to become sensitized and develop memory cells. The antibodies are also soon eliminated from her body. As a result, if she becomes pregnant again, her blood does not contain the anti-D that could harm the developing fetus.

Allergic reactions are directed against ordinary environmental antigens

About 20% of the U.S. population is plagued by an allergic disorder, such as allergic asthma or hay fever. A predisposition toward these disorders appears to be inherited. In allergic reactions, hypersensitivity results in the manufacture of antibodies against mild antigens, called allergens, which normally do not stimulate an immune response. Common environmental agents, such as pollen, house-dust mites, roach feces, and pet dander, can trigger allergic reactions in some individuals. In many kinds of allergic reactions, distinctive IgE immunoglobulins are produced. Let us examine a common allergic reaction, allergic rhinitis, commonly known as hay fever. This allergic response is triggered in some individuals by breathing pollen (FIG. 45-17). Exposure to pollen causes sensitization. Macrophages degrade the allergen and present fragments of it to T cells. The activated T cells then stimulate B cells to become plasma cells and produce pollen-specific IgE. These antibodies attach to receptors on mast cells. The Fc end of each IgE molecule Subsequent attaches to a mast cell receptor; its V First pregnancy pregnancy region is free to combine with the polBlood vessel of len allergen. the mother When a sensitized, allergic person inhales the microscopic pollen, allergen molecules rapidly attach to the IgE on sensitized mast cells. In response to IgE antibody binding with the allergen, mast cells release granules filled with histamine and other molecules that stimulate inflammation. Blood vessels dilate, and capillarPlacenta ies become more permeable, leading to edema and redness. Such responses + 1 Some Rh (antigen 2 Mother produces 3 Anti-D antibodies cause the nasal passages to become D–bearing) RBCs leak anti-D antibodies in cross placenta and swollen and irritated. A runny nose, across placenta from response to D antigens enter blood of fetus. fetus into mother’s blood. on Rh+ RBCs. Hemolysis of Rh+ sneezing, watery eyes, and a general blood occurs. feeling of discomfort are common. – RBC of mother Rh A prolonged allergic response oc+ Rh RBC of fetus with antigen D on surface curs when neutrophils are attracted by Anti-D antibody made against Rh+ RBC chemical compounds released by the Hemolysis of Rh+ RBC mast cells. Neutrophils migrate to the inflamed area and release compounds that lengthen the allergic reaction. In allergic asthma, an allergen– FIGURE 45-16 Animated Rh incompatibility IgE response occurs in the bronWhen an Rh-negative woman produces Rh-positive offspring, her immune system may become sensitized. chioles of the lungs. Mast cells release In subsequent pregnancies she may produce anti-D antibodies that cross the placenta and destroy fetal red blood cells. substances that cause smooth muscle

Nasal mucosa

Pollen grains 1

Previous exposure to pollen causes plasma cells to make pollen-specific IgE.

2

IgE combines with mast cell receptors in lining of nasal passages.

3

Pollen is inhaled.

4

Allergen combines with region of IgE on surface of sensitized mast cell.

5

Mast cell releases histamine and other chemicals.

6

This release causes increased vasodilation and increased capillary permeability. Symptoms result: edema, redness, constriction of respiratory passageways, mucus.

1

Plasma cell

IgE 3

2

Soluble antigens

Mast cell

4

5

Histamine and other compounds

6

Hay fever symptoms

FIGURE 45-17 An allergic reaction Many people respond to breathing pollen with allergic rhinitis (hay fever), a common type of allergic reaction.

to contract, and the airways in the lungs sometimes constrict for several hours, making breathing difficult. Allergic asthma is treated with steroids, which block cytokine production, and with drugs, such as leukotriene inhibitors, that block inflammatory mediators from binding to smooth muscle in the airway. Leukotrienes are inflammatory compounds released by mast cells. Leukotriene inhibitors reduce inflammation and decrease the incidence of asthma and hay fever episodes in individuals with hypersensitive immune systems. Monoclonal antibodies to IgE are also useful in treating these disorders because they reduce IgE levels in the blood. Certain foods or drugs act as allergens in some people, causing a reaction in the walls of the digestive tract that leads to discomfort and diarrhea. The allergen may be absorbed and cause mast cells to release granules elsewhere in the body. When the allergen–IgE reaction takes place in the skin, the histamine released by mast cells causes the swollen red welts known as urticaria, or hives. Systemic anaphylaxis is a dangerous allergic reaction that can occur when a person develops an allergy to a specific drug, such as penicillin; to compounds in the venom injected by a stinging insect; or even to certain foods. Within minutes after the substance enters the body, a widespread allergic reaction takes place. Mast cells release large amounts of histamine and other compounds into the circulation that cause extreme vasodilation and capillary permeability. So much plasma may be lost from the blood that circulatory shock and death can occur within a few minutes.

The symptoms of allergic reactions are often treated with antihistamines, drugs that block the effects of histamine. These drugs compete for the same receptors on cells targeted by histamine. When the antihistamine combines with the receptor, it prevents the histamine from binding and thus prevents its harmful effects. Antihistamines are useful clinically in relieving the symptoms of hives and hay fever. They are not completely effective because mast cells release molecules other than histamine that also cause allergic symptoms.

Graft rejection is an immune response against transplanted tissue Skin can be successfully transplanted from one part of the same body to another or from one identical twin to another. However, when skin is taken from one person and grafted onto the body of a nontwin, it is rejected and sloughs off. Why? Recall that tissues from the same individual or from identical twins have identical MHC alleles and thus the same MHC antigens. Such tissues are compatible. Because there are many alleles for each of the MHC genes, it is difficult to find identical matches among nonidentical twins. When a tissue or organ is taken from a donor and transplanted to the body of a nontwin host, several of the MHC antigens are likely to be different. The host’s immune system regards the graft

as foreign and launches an immune response known as graft rejection. In the first stage of rejection, the sensitization stage, TH cells and TC cells recognize the foreign antigens. In the second stage, the effector stage, TC cells, the complement system, and cytokines secreted by TH cells attack the transplanted tissue and can destroy it within a week. The sequence of events in graft rejection can be summarized as follows: tissue from donor transplanted into body of recipient ¡ TH cells and TC cells recognize MHC antigens on transplant cells as foreign ¡ TH cells secrete cytokines, complement system is activated, and TC cells destroy transplant cells

To avoid graft rejection, tissues from the patient and from potential donors must be typed and matched as closely as possible. Cell typing is somewhat similar to blood typing but is more complex. If all the MHC antigens are matched, the graft has about a 95% chance of surviving the first year. Unfortunately, not many people are lucky enough to have an identical twin to supply spare parts, so perfect matches are difficult to find. Furthermore, some organs, such as the heart, cannot be spared. Most transplanted organs, therefore, come from unrelated donors, often from patients who have just died. To prevent graft rejection in less compatible matches, physicians use antirejection drugs such as cyclosporine to suppress the immune system. Cyclosporine prevents T cells from producing cytokines. Unfortunately, immunosuppression makes the transplant patient more vulnerable to pneumonia or other infections and increases the risk of certain types of cancer. After the first few months, immunosuppressant drug dosages may be reduced. Researchers are working to develop more effective immunosuppression techniques.

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In the United States, thousands of patients are in need of organ transplants. Because the number of human donors does not meet this need, investigators are developing techniques for culturing stem cells as replacement tissues. Some researchers are working on more effective methods for transplanting animal tissues and organs to humans. Challenges include risks of transmitting animal diseases to humans and graft rejection. Pigs have been used as donors, but destructive immune responses are launched against these transplanted organs. Because they are closely related, chimpanzees and other primates can be used as donors with lower risk of graft rejection. However, ethical concerns have limited such transplants. Researchers are developing methods for genetically engineering pigs and other animals so that they do not produce antigens that stimulate immune responses in human recipients. These animals could then be cloned and used as donors of hearts, kidneys, and other organs. Effective artificial organs are also being developed. The body has a few immunologically privileged locations where foreign tissue is accepted. For example, corneal transplants are highly successful because the cornea has almost no associated blood or lymphatic vessels and is thus out of reach of most lymphocytes. Furthermore, antigens in the corneal graft probably would not find their way into the circulatory system and therefore could not stimulate an immune response.

Review How does the body defend itself against cancer? How does HIV infect the body? How does it affect the body? What is an autoimmune disease? Give two examples. What is the sequence of immunological events that take place in a common type of allergic reaction, such as hay fever? What is the immunological basis for graft rejection?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

45.1 (page 964) 1 Distinguish between nonspecific and specific immune responses. Immune responses are the body’s defensive responses against pathogens (disease-causing organisms), toxins, and other harmful agents. In immune responses, the body recognizes foreign or dangerous macromolecules and responds to eliminate them. Nonspecific immune responses provide general and immediate protection against pathogens, some toxins and drugs, and cancer cells. ■ Specific immune responses target distinct antigens, molecules recognized as foreign or dangerous by cells of the immune system. The immune system produces antibodies, highly specific proteins that recognize and bind to specific antigens. Another defining feature of specific immunity is immunological memory, that is, the system “remembers” foreign or dangerous molecules and responds more strongly to repeated encounters with the same molecules. 2 Compare, in general terms, the immune responses that have evolved in invertebrates and vertebrates. Invertebrates depend mainly on nonspecific immune responses, ■ such as physical barriers; phagocytosis (engulfing and destroying pathogens); receptors that recognize molecular patterns associated ■

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with pathogens; and antimicrobial peptides (soluble molecules that destroy pathogens). Vertebrates use both nonspecific and specific immune responses.

45.2 (page 966) 3 Describe nonspecific immune responses, including physical barriers; recognition by pattern recognition receptors; actions of phagocytes and other cells; actions of cytokines and complement; and the inflammatory response. ■ Nonspecific immune responses include physical barriers, such as the skin and the epithelial linings of the respiratory and digestive tracts. Antimicrobial peptides are produced by epithelial membranes. ■ When pathogens break through the first-line defenses, pathogenassociated molecular patterns, or PAMPs, bind with pattern recognition receptors and activate other nonspecific defenses. PAMPs are sets of common molecular features that are characteristic of each class of pathogens. The best known pattern recognition receptors are Toll-like receptors, receptors on phagocytes and certain other types of cells.

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Phagocytes, including neutrophils and macrophages, destroy bacteria. Natural killer (NK) cells destroy cells infected with viruses and foreign or altered cells such as tumor cells. Dendritic cells release antiviral proteins called interferons. Cytokines are signaling proteins that regulate interactions between cells. Important groups are interferons, tumor necrosis factors, interleukins, and chemokines. Interferons inhibit viral replication and activate natural killer cells. Tumor necrosis factor (TNF) stimulates immune cells to initiate an inflammatory response. Interleukins help regulate interactions between lymphocytes and other cells of the body, help mediate inflammation, and can mediate fever. Chemokines attract, activate, and direct the movement of certain cells of the immune system. Complement proteins lyse the cell wall of pathogens; coat pathogens, thereby enhancing phagocytosis; and attract white blood cells to the site of infection. These actions enhance the inflammatory response. When pathogens invade tissues, they trigger an inflammatory response, which includes three main processes: vasodilation, which brings more blood to the area of infection; increased capillary permeability, which allows fluid and antibodies to leave the circulation and enter the tissues; and increased phagocytosis. In response to infection, mast cells release histamine and other compounds that cause vasodilation and increased capillary permeability. Watch immune responses, including the inflammatory response, by clicking on the figures in CengageNOW.

45.4 (page 973) 6 Describe the sequence of events in cell-mediated immunity. ■

45.5 (page 975) 7 Summarize the sequence of events in antibody-mediated immunity, including the effects of antigen–antibody complexes on pathogens. ■ In antibody-mediated immunity, B cells are activated when they combine with antigen. Activation requires both an APC (such as a dendritic cell or macrophage) that has a foreign antigen– MHC complex displayed on its surface and a TH cell that secretes interleukins. ■ When activated B cells multiply, they give rise to clones of cells. The cloned cells differentiate and form plasma cells. Plasma cells produce specific antibodies, called immunoglobulins (Ig), in response to the specific antigens that activated them. An antibody combines with a specific antigen to form an antigen–antibody complex, which may inactivate the pathogen, stimulate phagocytosis, or activate the complement system. See cell-mediated and antibody-mediated immunity by clicking on the figures in CengageNOW.

45.3 (page 970) 4 Contrast cell-mediated and antibody-mediated immunity, and give an overview of each process. In cell-mediated immunity, specific T cells are activated; T cells release proteins that destroy cells infected with viruses or other intracellular pathogens. ■ In antibody-mediated immunity, specific B cells are activated; B cells multiply and differentiate into plasma cells, which produce antibodies. 5 Describe the functions of the principal cells of the specific immune system and of the major histocompatibility complex. ■ Two main types of cells important in specific immune responses are lymphocytes and antigen-presenting cells. Lymphocytes develop from stem cells in the bone marrow. T cells are lymphocytes responsible for cell-mediated immunity. The thymus gland confers immunocompetence on T cells by making them capable of distinguishing between self and nonself. There are several types of T cells, including T cytotoxic cells (TC cells), T helper cells (TH), memory TC cells, memory TH cells, and regulatory T cells (Tregs). T cells are distinguished by their T-cell receptors (TCRs). B cells are lymphocytes responsible for antibody-mediated immu■ nity. B cells differentiate into plasma cells, which produce antibodies. Some activated B cells become memory B cells, which continue to produce antibodies after the body overcomes the infection. ■ Antigen-presenting cells (APCs), including dendritic cells, macrophages, and B cells, display foreign antigens as well as their own surface proteins. Dendritic cells are specialized to process, transport, and present antigens. ■ Immune responses depend on the major histocompatibility complex (MHC), a group of genes that encode MHC proteins. Class I MHC genes encode self-antigens, glycoproteins expressed on the surface of most nucleated cells. Class II MHC genes encode glycoproteins expressed on APCs of the immune system. Class III MHC genes encode components of the complement system and TNFs. ■

In cell-mediated immunity, specific T cells are activated by a foreign antigen–MHC complex on the surface of an infected cell. A costimulatory molecule and interleukins are also required. Activated TC cells multiply and give rise to a clone, a process called clonal expansion. These differentiated cells migrate to the site of infection and destroy pathogen-infected cells. Activated TH cells give rise to a clone of TH cells, which secrete cytokines that activate B cells and macrophages.

8 Describe the basic structure and function of an antibody, and explain the basis of antibody diversity. ■ An antibody molecule consists of four polypeptide chains: two identical heavy chains and two shorter light chains. Each chain has a constant (C) region and a variable (V) region. In a Y-shaped antibody, the two arms combine with antigen.

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Rearrangement of DNA segments during the differentiation of B cells is the main factor responsible for antibody diversity; millions of different types of B (and T) cells are produced. Learn more about antibody structure, function, and diversity by clicking on the figures in CengageNOW.

9 Describe the basis of immunological memory, and contrast secondary and primary immune responses. ■ After an infection, memory B cells and memory T cells remain in the body. These cells are responsible for long-term immunity. ■ The first exposure to an antigen stimulates a primary immune response. A second exposure to the same antigen evokes a secondary immune response, which is more rapid and more intense than the primary response.

10 Compare active and passive immunity, and give examples of each. ■

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Active immunity develops as a result of exposure to antigens; it may occur naturally after recovery from a disease or can be artificially induced by immunization with a vaccine. Passive immunity is a temporary condition that develops when an individual receives antibodies produced by another person or animal.

45.6 (page 982) 11 Describe the body’s response to cancer cells and HIV. ■

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NK cells, macrophages, T cells, and other cells of the immune system recognize antigens on cancer cells and launch an immune response against them. Cancer cells evade the immune system by blocking TC directly or by decreasing the class I MHC molecules on TC cells. Acquired immunodeficiency syndrome (AIDS) is caused by the human immunodeficiency virus (HIV), a retrovirus. HIV destroys

T helper cells, severely impairing immunity and placing the patient at risk for opportunistic infections. 12 Describe examples of hypersensitivity, including autoimmune diseases, Rh incompatibility, and allergic reactions, and summarize the immunological basis of graft rejection. ■ In autoimmune diseases, the body reacts immunologically against its own tissues. When an Rh-negative woman gives birth to an Rh-positive baby, she ■ may develop anti-D antibodies. Rh incompatibility can then occur in future pregnancies. ■ In an allergic reaction, an allergen stimulates the production of IgE, which combines with receptors on mast cells. The mast cells release histamine and other molecules that cause inflammation and other symptoms of allergy. Systemic anaphylaxis is a rapid, widespread allergic reaction that can lead to death. ■ Transplanted tissues have MHC antigens that stimulate graft rejection, an immune response in which T cells destroy the transplant.

T E S T YO U R U N D E R S TA N D I N G 1. Nonspecific (innate) immune responses include all but (a) inflammation (b) antigen–antibody complexes (c) binding of PAMPs with pattern recognition receptors (d) complement (e) phagocytosis

9. Immunoglobulin A (a) recognizes pathogen-associated molecular patterns (b) binds with NK cells (c) prevents pathogens from attaching to epithelial surfaces (d) is found mainly on dendritic cell surfaces (e) is found mainly on T-cell surfaces

2. Cytokines (a) are regulatory Toll-like receptors (b) prevent the inflammatory response (c) include interferons and interleukins (d) are immunoglobulins (e) b and c are correct

10. When a person is exposed to the same antigen a second time, the response is (a) less specific (b) more rapid (c) mediated by dendritic cells (d) likely to lead to development of an autoimmune disease (e) a hypersensitivity reaction

3. Which of the following is not an action of complement? (a) enhances phagocytosis (b) enhances inflammatory response (c) coats pathogens (d) lyses viruses (e) stimulates production of memory T cells 4. Which of the following cells are antigen-presenting cells? (a) NK cells and monocytes (b) macrophages and plasma cells (c) dendritic cells and macrophages (d) mast cells and B cells (e) memory T cells and plasma cells

11. In an allergic reaction, (a) an allergen binds with IgE (b) the body is immunodeficient (c) T helper cells release histamine (d) allergens are produced by plasma cells (e) mast cells are deactivated 12. Label the diagram and explain what this diagram illustrates. (Consult Figure 45-7, step 3 to check your answers.)

5. Which of the following cells are especially adept at destroying tumor cells? (a) NK cells (b) plasma cells (c) neutrophils (d) B cytotoxic cells (e) mast cells 6. Which of the following cells become immunologically competent after processing in the thymus gland? (a) NK cells (b) T cells (c) macrophages (d) B cells (e) plasma cells 7. The major histocompatibility complex (MHC) (a) consists of Y-shaped molecules (b) encodes certain antibodies (c) encodes Toll-like receptors (d) inhibits complement release from macrophages (e) encodes a group of cell-surface proteins 8. Which sequence most accurately describes antibody-mediated immunity? 1. B cell divides and gives rise to clone 2. antibodies produced 3. cells differentiate and form plasma cells 4. activated T helper cell interacts with B cell displaying same antigen complex 5. B cell activated (a) 1, 2, 3, 4, 5 (b) 3, 2, 1, 4, 5 (c) 4, 5, 3, 2, 1 (d) 4, 5, 1, 3, 2 (e) 4, 3, 1, 2, 5

TH cell

Activated B cell

Cell division

CRITICAL THINKING 1. Specificity, diversity, and memory are key features of the immune system. Giving specific examples, explain how each of these features is important. 2. What would be the effects of total loss of macrophages in the body? Which do you think would have a greater effect on the immune system, loss of macrophages or loss of B cells? 3. What are the advantages of having MHC antigens? disadvantages? What do you think would be the consequences of not having them? 4. A playmate in kindergarten exposes John and Jack to measles. John has been immunized against measles, but Jack has not received the measles vaccine. How are their immune responses different? Five years later, John and Jack are playing together when Judy, who is coming down with measles, sneezes on both of them. Compare their immune responses. 5. EVOLUTION LINK. The enzymes Rag1 and Rag2 cut and rearrange DNA, producing millions of specific antibodies and T-cell receptors. Until recently, immunologists thought that the genes

(Rag1 and Rag2) that encode these enzymes were found only in jawed vertebrates. However, investigators recently found similar genes in the purple sea urchin. What does this finding suggest about the evolution of the specific immune system? 6. EVOLUTION LINK. Fishes have only one class of antibodies, IgM. Amphibians have two classes, and the number increases to five main classes in mammals. Hypothesize an adaptive advantage of having a greater variety of antibody groups. 7. SCIENCE, TECHNOLOGY, AND SOCIETY. Imagine that you are a researcher developing new HIV treatments. What approaches might you take? What public policy decisions would you recommend that might help slow the spread of AIDS while new treatments or vaccines are being developed? Additional questions are available in CengageNOW at www.cengage.com/ login.

46

Gas Exchange

Skip Moody/Dembinsky Photo Associates

Ventilation. Terrestrial vertebrates, like the whitetailed deer (Odocoileus virginanus), ventilate their lungs by breathing air.

KEY CONCEPTS

M

46.1 Air has a higher concentration of molecular oxygen

damaged beyond repair if their oxygen supply is cut off for only a few

than water does, and animals require less energy to move air than to move water over a gas exchange surface; however, animals that respire in air must have adaptations that reduce water loss so that their respiratory surfaces do not dry out.

minutes. To stay alive, animals must take in oxygen and release excess

46.2 Adaptations for gas exchange include a thin, moist

oxygen from the environment is taken up by the animal and delivered to

body surface; gills in aquatic animals; and tracheal tubes and lungs in terrestrial animals.

its individual cells. At the same time, carbon dioxide generated during

46.3 In mammals, oxygen and carbon dioxide are ex-

respiration takes place in mitochondria. Oxygen is essential because it

changed between alveoli and blood by diffusion; the pressure of a particular gas determines its direction and rate of diffusion. Respiratory pigments combine with oxygen and transport it; carbon dioxide is transported mainly as bicarbonate ions.

46.4 Several defense mechanisms protect the lungs from harmful substances that are inhaled; tobacco smoke and other air pollutants are major causes of disease, disability, and death.

ost animal cells require a continuous supply of oxygen for cellular respiration. Mammalian brain cells, and some other cells, may be

carbon dioxide. The exchange of gases between an organism and its environment is called respiration. There are two phases of respiration. During organismic respiration,

cellular respiration is excreted into the environment. Aerobic cellular serves as the final electron acceptor in the mitochondrial electron transport chain (see Chapter 8). Carbon dioxide is a metabolic waste product of cellular respiration. Sponges, hydras, flatworms, and many other small, aquatic organisms exchange gases entirely by simple diffusion, the passive movement of particles (atoms, ions, or molecules) from a region of higher concentration to a region of lower concentration, that is, down a concentration

gradient (see Chapter 5). In these animals, most cells are in direct contact with the environment. Dissolved oxygen from the surrounding water diffuses into the cells, while carbon dioxide diffuses

again pass through these airways (providing opportunity for retaining water) before leaving the body. These adaptations protect the lungs from the drying and cooling effects of air.

out of the cells and into the water. No specialized respiratory structures are needed. In an animal more than about 1 mm thick, oxygen cannot dif-

Review ■

fuse quickly enough through layers of cells to support life. Special-

What are some advantages of gas exchange in air over gas exchange in water?

ized respiratory structures, such as gills or lungs, deliver oxygen to the cells or to a transport system. These structures also facilitate excretion of carbon dioxide. Animals with lungs carry on ventilation by breathing air (see photograph). In many animal groups, including vertebrates, respiratory

46.2 TYPES OF RESPIRATORY SURFACES

systems and circulatory systems are functionally connected. They

■ ■ LEARNING OBJECTIVE

work together to provide efficient intake and transport of oxygen.

2

Such systems provide sufficient oxygen to support high metabolic rates. As we will discuss, different strategies for gas exchange have evolved in the arthropods. In this chapter, we will examine adaptations for gas exchange in air and water and describe various types of respiratory surfaces. We will then focus on the structure and function of the mammalian respiratory system. We end the chapter with a discussion of the effects of breathing polluted air, including cigarette smoke.

46.1 ADAPTATIONS FOR GAS EXCHANGE IN AIR OR WATER ■ ■ LEARNING OBJECTIVE 1

Compare the advantages and disadvantages of air and water as mediums for gas exchange, and describe adaptations for gas exchange in air.

Gas exchange in air has certain advantages over gas exchange in water. Air contains a much higher concentration of molecular oxygen (at least 20 times more) than water does. In addition, oxygen diffuses about 10,000 times faster through air than through water. Another advantage is that less energy is needed to move air than to move water over a gas exchange surface because air is less dense and less viscous. A mammal uses less than 2% of its energy budget to breathe compared to a fish, which must spend as much as 20% of its energy to obtain its oxygen from water. Gas exchange in air presents a major challenge: the threat of desiccation. Animals that respire in air struggle continuously with water loss. Respiratory surfaces must be kept moist because oxygen and carbon dioxide are dissolved in the fluid that bathes the cells of these surfaces. Whether an animal makes its home on land or water, gas exchange takes place across a moist surface. Adaptations have evolved that keep respiratory surfaces moist and minimize desiccation. For example, the lungs of air-breathing vertebrates are located deep within the body, not exposed as gills are. Air is humidified and brought to body temperature as it passes through the moist upper respiratory passageways. Expired air must

Describe the following adaptations for gas exchange: body surface, tracheal tubes, gills, and lungs.

Respiratory structures must be moist, and must also have thin walls through which diffusion can easily occur. Four main types of respiratory surfaces have evolved in animals: the animal’s own body surface, tracheal tubes, gills, and lungs (FIG. 46-1). Some animals use a combination of these adaptations. Gills are adapted for gas exchange in water, and the tracheal tubes of insects and lungs of vertebrates are respiratory structures adapted for gas exchange in air. With the important exception of insect tracheal tubes, respiratory structures are richly supplied with blood vessels that facilitate exchange and transport of respiratory gases. Most animals carry on ventilation, that is, they actively move air or water over their respiratory surfaces. If the air or water supplying oxygen to the cells can be continuously renewed, more oxygen will be available. Sponges use their flagella to set up a current of water through the channels of their bodies. Most fishes gulp water, which then passes over their gills. Terrestrial vertebrates have lungs and breathe air. When mammals breathe, the diaphragm and other muscles move air in and out of the lungs.

The body surface may be adapted for gas exchange Gas exchange occurs through the entire body surface in many animals, including nudibranch mollusks, most annelids, and some amphibians. These animals are small, with a high surface area to volume ratio. They also have a low metabolic rate that requires smaller quantities of oxygen per cell. In aquatic animals, the body surface is kept moist by the surrounding water. In terrestrial animals, the body secretes fluids that keep its surface moist. Many animals that exchange gases across the body surface also have gills or lungs.

Tracheal tube systems deliver air directly to the cells In insects and some other arthropods (such as chilopods, diplopods, some mites, and some spiders), the respiratory system is a network of tracheal tubes, also called tracheae (FIG. 46-2). This very efficient respiratory system delivers air directly to the cells.

KEY POINT

The efficient tracheal tube system of insects delivers air directly to the cells.

Earthworm

(a) Body surface. Some small, multicellular animals exchange gases through the body surface. Grasshopper

Spiracle

(b) Tracheal tubes. Insects and some other arthropods exchange gases through a system of tracheal tubes, or tracheae.

Tracheal tube

(a) Location of spiracles and tracheal tubes. Air enters the system of tracheal tubes through openings called spiracles. Epithelial cell

Internal gills O2

Tracheal tube

External gills

Spiracle

Tracheole

CO2

Gills Mud puppy

Fish

(c) Gills. Most aquatic animals exchange gases through gills, thin structures that extend from the body surface. Gills can be internal or external.

Muscle

Book lung

Lungfish Spider

Mammal

Courtesy of Dr. James L. Nation and Stain Technology, Vol. 58, 1983

(b) Structure and function of tracheal tubes. Air passes through a system of branching tracheal tubes that conduct oxygen to all cells of the insect.

100 μm

(c) SEM of a mole cricket trachea. The wrinkles are a part of a long spiral that wraps around the tube, strengthening the tracheal wall somewhat as a spring strengthens the plastic hoses of many vacuum cleaners. The tracheal wall is composed of chitin.

(d) Lungs. Lungs are adaptations for terrestrial gas exchange.

FIGURE 46-1 Adaptations for gas exchange

FIGURE 46-2 Gas exchange in insects: tracheal tubes

Air enters the tracheal tubes through a series of as many as 20 tiny openings called spiracles along the body surface. In some insects, especially large, active ones, muscles help ventilate the tracheae by pumping air in and out of the spiracles. For example, the grasshopper draws air in through the first four pairs of spiracles when the abdomen expands. Then the abdomen contracts, forcing air out through the last six pairs of spiracles. Once inside the body, the air passes through a system of branching tracheal tubes, which extend to all parts of the animal. The tracheal tubes terminate in microscopic, fluid-filled tracheoles. Gases are exchanged between this fluid and the body cells. The tracheal system supplies enough oxygen to support the high metabolic rates required by many insects.

Gills are the respiratory surfaces in many aquatic animals Found mainly in aquatic animals, gills are moist, thin structures that extend from the body surface. They are supported by the buoyancy of water but tend to collapse in air. In many animals, the outer surface of the gills is exposed to water, whereas the inner side is in close contact with networks of blood vessels. Sea stars and sea urchins have dermal gills that project from the body wall. Their ciliated epidermal cells ventilate the gills by beating a stream of water over them. Gases are exchanged between the water and the coelomic fluid inside the body by diffusion through the gills. Various types of gills are found in some annelids, aquatic mollusks, crustaceans, fishes, and amphibians. Mollusk gills are folded, providing a large surface for respiration. In clams and other bivalve mollusks and in simple chordates, gills may also be adapted for trapping and sorting food. The rhythmic beating of cilia draws water over the gill area, and food is filtered out of the water while gases are exchanged. In mollusks, gas exchange also takes place through the mantle. In chordates, gills are usually internal. A series of slits perforates the pharynx, and the gills lie along the edges of these gill slits (see Fig. 32-4). In bony fishes, the fragile gills are protected by an external bony plate, the operculum. In some fishes, movements of the jaw and operculum help pump water rich in oxygen through the mouth and across the gills. The water leaves through the gill slits. Each gill in the bony fish consists of many gill filaments, which provide an extensive surface for gas exchange (FIGS. 46-3a and b). The filaments extend out into the water, which continuously flows over them. A capillary network delivers blood to the gill filaments, facilitating diffusion of oxygen and carbon dioxide between blood and water. This system is extremely efficient because blood flows in a direction opposite to the movement of the water. This arrangement, called a countercurrent exchange system, maximizes the difference in oxygen concentration between blood and water throughout the area where the two remain in contact (FIG. 46-3c). If blood and water flowed in the same direction, that is, concurrent exchange, the difference between the oxygen concentrations in blood (low) and water (high) would be very large initially and very small at the end (FIG. 46-3d). The oxygen concentration in the

water would decrease as the concentration in the blood increased. When the concentrations in the two fluids became equal, equilibrium would be reached, and the net diffusion of oxygen would stop. Only about 50% of the oxygen dissolved in the water could diffuse into the blood. In the countercurrent exchange system, however, blood low in oxygen comes in contact with water that is partly depleted of oxygen. Then, as the blood flowing through the capillaries becomes progressively richer in oxygen, the blood comes in contact with water with an increasingly higher concentration of oxygen. Thus, all along the capillaries, the diffusion gradient favors passage of oxygen from the water into the gill. A high rate of diffusion is maintained, ensuring that a very high percentage (more than 80%) of the available oxygen in the water diffuses into the blood. Oxygen and carbon dioxide do not interfere with each other’s diffusion, and they simultaneously diffuse in opposite directions. The reason is that oxygen is more concentrated outside the gills than inside but carbon dioxide is more concentrated inside the gills than outside. Thus, the same countercurrent exchange mechanism that ensures efficient inflow of oxygen also results in equally efficient outflow of carbon dioxide.

Terrestrial vertebrates exchange gases through lungs Lungs are respiratory structures that develop as ingrowths of the body surface or from the wall of a body cavity such as the pharynx (throat region). For example, the book lungs of spiders are enclosed in an inpocketing of the abdominal wall. These lungs consist of a series of thin, parallel plates of tissue (like the pages of a book) filled with hemolymph (see Fig. 46-1d). The plates of tissue are separated by air spaces that receive oxygen from the outside environment through a spiracle. A different type of lung evolved in land snails and slugs. (These terrestrial mollusks lack gills.) Gas exchange takes place through a lung, which is a vascular region of the mantle. Fossil evidence suggests that early lobe-finned fishes had lungs somewhat similar to those of modern lungfishes. The Australian lungfish can use either its gills or its lungs, depending on the conditions in its environment. The gills of African and South American lungfishes degenerate with age, so adults must rise to the surface and exchange gases entirely through their lungs. Some paleontologists hypothesize that all early bony fishes had lungs or lunglike structures. Most modern bony fishes have no lungs, but nearly all have homologous swim bladders (see Chapter 32). By adjusting the amount of gas in its swim bladder, the fish can control its buoyancy. Most amphibians have lungs (FIG. 46-4). However, most of their gas exchange takes place across the body surface. The lungs of salamanders are two long, simple sacs richly supplied with capillaries. Frogs and toads have ridges containing connective tissue on the inside of the lungs, which increases the respiratory surface. Some amphibians do not have lungs. Among plethodontid (lungless) salamanders, for example, gas exchange takes place across the thin, wet skin and through the moist, vascular walls of the mouth and pharynx.

KEY POINT

Countercurrent flow efficiently oxygenates blood circulating through the gill filaments.

Gill arch

CO2

Flow of H2O Opercular chamber

(d) Concurrent flow (hypothetical). If the system were concurrent, that is, if blood flowed through the capillaries in the same direction as the flow of the water, much less of the oxygen dissolved in the water could diffuse into the blood.

(a) Location of gills. The gills form the lateral wall of the pharyngeal cavity. They lie under a bony plate, the operculum, which has been removed in this side view. Gill arch Blood vessels Gill filaments

Afferent blood vessel (low O2 concentration)

Efferent blood vessel (rich in O2)

Bernard Photo Production / Animals Animals

(b) Structure of a gill. Each gill consists of a cartilaginous gill arch to which two rows of leaflike gill filaments attach. Each gill filament has many smaller extensions rich in capillaries. As water flows past the gill filament, oxygen passes from the water into blood circulating through the capillaries.

(e) Gills of the salmon.

(c) Countercurrent flow. Blood entering the capillaries of the gill filament is deficient in oxygen. The blood flows through the capillaries in a direction opposite to that taken by the water. This countercurrent exchange system efficiently charges the blood with oxygen.

FIGURE 46-3 Animated Gas exchange in bony fishes: gills

Reptiles have better developed lungs than amphibians. The respiratory surface of each lung is divided into large sacs that increase the surface for gas exchange. In some turtles, lizards, and crocodiles, the lungs have many subdivisions that give them a spongy texture. Reptiles ventilate their lungs by drawing air into the lungs,

but the muscles involved differ in the various reptile groups. Gas exchange is not very efficient and does not supply enough oxygen to sustain long periods of activity. Birds have the most efficient respiratory system of any living vertebrate. Very active, endothermic animals with high metabolic

KEY POINT

During vertebrate evolution, the surface area of the lung increased.

Trachea To other lung

Salamander's lungs

Frog's lungs

Toad's lung

Trachea

flows through them. Gas exchange takes place across the walls of these tubes, which are rich in capillaries. Chickens and other weakflying birds have about 400 parabronchi per lung; pigeons and other strong-flying birds have about 1800 parabronchi per lung. The direction of blood flow in the lungs is in a different direction from that of airflow through the parabronchi. This arrangement, similar in principle to the countercurrent exchange in the gills of fishes, increases the amount of oxygen that enters the blood. However, in birds the capillaries are oriented at right angles to the parabronchi rather than along their length. For this reason, this arrangement is referred to as crosscurrent rather than countercurrent.

Review ■

■

To other lung

■

Why are specialized respiratory structures necessary in a tadpole but not in a flatworm? How does gas exchange differ among the following animals: (1) earthworm, (2) grasshopper, (3) fish, and (4) bird? How does the countercurrent exchange system increase the efficiency of gas exchange between a fish’s gills and blood?

Air sac

Air sac Reptile's lung

Bird's lungs

FIGURE 46-4 Animated Evolution of vertebrate lungs Salamander lungs are simple sacs. Other amphibians and reptiles have lungs with small ridges or folds that increase surface area. Birds have an elaborate system of lungs and air sacs. Mammalian lungs have millions of air sacs (alveoli) that increase the surface available for gas exchange (see Fig. 46-7).

46.3 THE MAMMALIAN RESPIRATORY SYSTEM ■ ■ LEARNING OBJECTIVES 3 4 5

6

rates, birds require large amounts of oxygen to sustain flight and other activities. Their small, bright red lungs have extensions (usually nine) called air sacs, which reach into all parts of the body and even connect with air spaces in some of the bones. The air sacs act as bellows, drawing air into the system. Collapse of the air sacs during exhalation forces air out. Gas exchange does not take place across the walls of the air sacs. The high-performance bird respiratory system is arranged so that air flows in one direction through the lungs and is renewed during a two-cycle process (FIG. 46-5). Air entering the body passes into the posterior air sacs and the part of the lungs closest to these air sacs. The posterior air sacs are reservoirs for fresh air. When the bird exhales, that air flows into the lungs. At the second breath, the air flows from the lungs to the anterior air sacs. Finally, at the second exhalation, the air leaves the body as another breath of air enters the lungs. Thus, a bird gets fresh air across its lungs through both inhalation and exhalation. In birds the narrowest branches of the airway end in tiny, thinwalled tubes called parabronchi. In contrast to the saclike alveoli of mammals, parabronchi are open at both ends and air continuously

Trace the passage of oxygen through the human respiratory system from nostrils to alveoli. Summarize the mechanics of breathing in humans, and describe gas exchange in the lungs and tissues. Summarize the mechanisms by which oxygen and carbon dioxide are transported in the blood and identify factors that determine the oxygen–hemoglobin dissociation curve. Describe the regulation of breathing in humans and summarize the physiological effects of hyperventilation and of sudden decompression when a diver surfaces too quickly from deep water.

The respiratory system of mammals consists of the lungs and a series of tubes through which air passes on its journey from the nostrils to the lungs and back (FIG. 46-6). The complex lungs have an enormous surface area. As in all vertebrates, the respiratory and circulatory systems are functionally connected. The circulatory system delivers oxygen-rich blood to all of the cells in the body. In the following sections, we focus mainly on the human respiratory system.

The airway conducts air into the lungs A breath of air enters the body through the nostrils and flows through the nasal cavities. Air passing through the nose is filtered, moistened, and brought to body temperature. The nasal cavities are lined with a moist, ciliated epithelium rich in blood vessels. Inhaled dirt, bacteria, and other foreign particles are trapped in the stream of mucus produced by cells within the epithelium and pushed along toward the throat by the cilia. In this way, foreign particles are de-

Trachea Air sacs Air Lung

Anterior air sacs

Posterior air sacs

(a) Structure of the bird respiratory system.

(b) First inhalation. As the bird inhales, fresh air (green) flows into the posterior air sacs and partly into the lungs (not shown).

(c) First exhalation. As the bird exhales, air from the posterior air sacs is forced into the lungs.

(d) Second inhalation. Air from the first breath moves into the anterior air sacs and partly into the lungs (not shown). Air from the second inhalation (gray) flows into the posterior air sacs.

(e) Second exhalation. Most of the air from the first inhalation leaves the body, and air from the second inhalation flows into the lungs.

FIGURE 46-5 Animated Gas exchange in birds The bird respiratory system includes lungs and air sacs. The bird’s breathing process requires two cycles of inhalation and exhalation to support a one-way flow of air through the lungs.

Respiratory centers

Sinuses Nasal cavity Tongue

Pharynx

Epiglottis Larynx

Esophagus

Space occupied by heart

Trachea

Bronchioles

livered to the digestive system, which can more effectively dispose of such materials than can the delicate lungs. A person normally swallows more than a pint of nasal mucus each day, and even more during an infection or allergic reaction. The back of the nasal cavities is continuous with the throat region, or pharynx. Air finds its way into the pharynx whether one breathes through the nose or mouth. An opening in the floor of the pharynx leads into the larynx. Because the larynx contains the vocal cords, it is also referred to as the “voice box.” Cartilage embedded in its wall prevents the larynx from collapsing and makes it hard to the touch when felt through the neck.

Diaphragm

Left lung

▲

Bronchus Right lung

FIGURE 46-6 Animated The human respiratory system The internal view of one lung illustrates a portion of its extensive system of air passageways. The muscular diaphragm forms the floor of the thoracic cavity. The respiratory centers in the brain regulate the rate of respiration.

KEY POINT

Oxygen diffuses through the thin wall of the alveolus and then through the thin walls of the surrounding capillaries to reach the blood.

Bronchiole

Alveolus

Capillary Macrophage Red blood cells Capillaries Epithelial cell of the adjacent alveolus

Alveolus

Epithelial cell of the wall of the alveolus

Alveolus

Dr. Richard Kessel & Dr. Randy K/ Visuals Unlimited/ Corbis

Courtesy of Drs. Peter Gehr, Marianne Bachofen, and Ewald R. Wiebel

(a) A cluster of alveoli. Gas exchange takes place across the thin wall of the alveolus. Extensive capillary networks lie between the walls of adjacent alveoli.

Wall of alveolus

Wall of capillary

Red blood cell

1 μm

(c) Color-enhanced TEM of a portion of a capillary and the wall of an alveolus. The dark structure extending through the capillary is part of a red blood cell. Notice the very short distance oxygen must diffuse to get from the air within the alveolus to the red blood cells that transport it to the body tissues. 30 μm

(b) Capillaries in lung. Capillaries surround the individual alveoli (air sacs), allowing for the exchange of oxygen and carbon dioxide between the blood and the alveoli.

FIGURE 46-7 Animated Structure and function of alveoli The alveoli are the air sacs through which gases are exchanged between the air in the lung and the blood in the capillaries.

During swallowing, a flap of tissue called the epiglottis automatically closes off the larynx so that food and liquid enter the esophagus rather than the lower airway. If this mechanism fails and foreign matter enters the sensitive larynx, a cough reflex expels the material. Despite these mechanisms, choking sometimes occurs. From the larynx, air passes into the trachea, or windpipe, which is kept from collapsing by rings of cartilage in its wall. The trachea divides into two branches, the bronchi (sing., bronchus); one bronchus connects to each lung. Both trachea and bronchi are lined by a mucous membrane containing ciliated cells. These cells trap medium-sized particles that escape the cleansing mechanisms of the nose and larynx. Mucus containing these particles is constantly beaten upward by the cilia to the pharynx, where it is periodically swallowed. This mechanism, functioning as a ciliapropelled elevator of mucus, helps keep foreign material out of the lungs.

Gas exchange occurs in the alveoli of the lungs The lungs are large, paired, spongy organs occupying the thoracic (chest) cavity. The right lung is divided into three lobes, the left lung into two lobes. Each lung is covered with a pleural membrane, which forms a continuous sac that encloses the lung and extends outward to become the lining of the thoracic cavity. The pleural cavity is the space between the pleural membranes. A film of fluid in the pleural cavity provides lubrication between the lungs and the chest wall. Because the lung consists largely of air tubes and elastic connective tissue, it is a spongy, elastic organ with a very large internal surface area for gas exchange. Within each lung the bronchi branch, becoming progressively shorter, narrower, and more numerous, similar to the branching of a tree. Small branches give rise to the even smaller bronchioles in each lung. Each bronchiole ends in a cluster of tiny air sacs, the alveoli (sing., alveolus) (FIG. 46-7). Each human lung contains more than 200 million alveoli, which provide an internal surface area the approximate size of a tennis court! Each alveolus is lined by an extremely thin, single layer of epithelial cells. Gases diffuse freely through the wall of the alveolus and into the capillaries that surround it. Only two thin cell layers—the epithelium of the alveolar wall and the epithelium of the capillary wall—separate the air in the alveolus from the blood. In summary, air passes through the following sequence of structures after it enters the body: nostrils ¡ nasal cavities ¡ pharynx ¡ larynx ¡ trachea ¡ bronchi ¡ bronchioles ¡ alveoli

Ventilation is accomplished by breathing Breathing is the mechanical process of moving air from the environment into the lungs and of expelling air from the lungs. Inhaling air is called inhalation or inspiration; exhaling air is exhalation or expiration. The thoracic cavity is closed so no air can enter except through the trachea. (When the chest wall is punctured, for example, by a fractured rib or gunshot wound, air enters the pleural space and the lung collapses.)

During inhalation, the volume of the thoracic cavity is increased by the contraction of the diaphragm, the dome-shaped muscle that forms its floor. When the diaphragm contracts, it moves downward, increasing the volume of the thoracic cavity (FIG. 46-8). During forced inhalation, when a large volume of air is inhaled, the external intercostal muscles contract as well. This action moves the ribs upward, which also increases the volume of the thoracic cavity. Because the lungs adhere to the walls of the thoracic cavity, when the volume of the thoracic cavity increases, the space within each lung also increases. The air in the lungs now has more space in which to move about. As a result, air pressure in the lungs falls by 2 or 3 millimeters of mercury (mm Hg) below the air pressure outside the body. Because of this pressure difference, air from the

KEY POINT

Changes in position of the diaphragm and of the external and internal intercostal muscles in inhalation and exhalation change the volume of the thoracic cavity.

Trachea Lung

Diaphragm

(a) Inhalation. The diaphragm contracts, increasing the volume of the thoracic cavity. Air moves into the lungs.

(b) Exhalation. The diaphragm relaxes, decreasing the volume of the thoracic cavity. Air moves out of the lungs.

Diaphragm

(c) Forced inhalation. The external intercostal muscles contract, pulling the rib cage upward and outward. This increases the front-to-back dimension of the chest and correspondingly increases the volume of the thoracic cavity.

(d) Forced exhalation. The internal intercostal muscles contract, pulling the rib cage downward and inward. The volume of the thoracic cavity decreases.

FIGURE 46-8 Animated Mechanics of breathing

outside rushes in through the respiratory passageways and fills the lungs until the two pressures are equal once again. Exhalation occurs when the diaphragm relaxes. The volume of the thoracic cavity decreases, raising the pressure in the lungs to 2 to 3 mm Hg above atmospheric pressure. The millions of distended air sacs partially deflate and expel the inhaled air. The pressure then returns to normal, and the lung is ready for another inhalation. To summarize: during inhalation the millions of alveoli fill with air like so many tiny balloons. Then, during exhalation, the air rushes out of the alveoli, partially deflating them. During deep or forced inhalation, the external intercostal muscles also contract, pulling the rib cage upward and outward. This action further increases the volume of the thoracic cavity and more air moves into the lungs. During forced exhalation, muscles of the abdominal wall and the internal intercostal muscles contract, pushing the diaphragm up and the ribs down. This action decreases the volume of the thoracic cavity, pushing air out of the lungs. Some of the work in stretching the thorax and the lungs is necessary to stretch elastic connective tissue. Work is also required to overcome the cohesive force of water molecules associated with the pleural membranes. The forces between the water molecules produce a surface tension that resists stretching. The work of breathing is reduced by pulmonary surfactant, a detergent-like phospholipid mixture secreted by specialized epithelial cells in the lining of the alveoli. Pulmonary surfactant intersperses between the water molecules, reducing their cohesive force. This action markedly reduces the surface tension of the water, prevents the alveoli from collapsing, and reduces the energy required to stretch the lungs. Premature infants often cannot produce enough surfactant, which causes respiratory distress syndrome. In these infants, high surface tension makes it difficult to inflate the lungs, and the alveoli collapse during expiration. Breathing is labored. Respiratory failure as a result of surfactant deficiency is a leading cause of morbidity and mortality in premature infants. Surfactant deficiency can be treated with surfactant replacement therapy in which artificial surfactant is delivered through the trachea.

Each alveolus serves as a tiny depot from which oxygen diffuses into the blood (FIG. 46-9). Oxygen molecules efficiently pass by simple diffusion from the alveoli, where they are more concentrated, into the blood in the pulmonary capillaries, where they are less concentrated. At the same time, carbon dioxide moves from the blood, where it is more concentrated, to the alveoli, where it is less concentrated. Each gas diffuses through the single layer of cells lining the alveoli and the single layer of cells lining the capillaries. Recall that in aerobic respiration, oxygen is the final electron acceptor; it accepts two electrons, which are added to two protons to produce water (see Chapter 8). Inhaled (atmospheric) air contains about 20.9% oxygen, but because oxygen is used during cellular respiration, exhaled (alveolar) air contains only 14% oxygen. Carbon dioxide is produced during aerobic respiration. For this reason, exhaled air contains more than 100 times as much (5.6%) carbon dioxide as inhaled air (almost .04% carbon dioxide). The concentration of oxygen in the cells is lower than in the capillaries entering the tissues, and the concentration of carbon dioxide is higher in the cells than in the capillaries. As blood circulates through capillaries of a tissue such as brain or muscle, oxygen moves by simple diffusion from the blood to the cells, and carbon dioxide moves from the cells into the blood. The factor that determines the direction and rate of diffusion is the pressure or tension of the particular gas. According to Dalton’s

PO = 100 mm Hg 2 PCO = 40 mm Hg 2

Alveoli in lung

(a) In the alveoli. The concentration of oxygen is greater than in the pulmonary capillaries, so oxygen diffuses from the alveoli into the blood. Carbon dioxide is more concentrated in the blood than in the alveoli, so it diffuses out of the O2 capillaries and into the CO2 alveoli.

Capillary in tissue

The quantity of respired air can be measured The amount of air moved into and out of the lungs with each normal resting breath is called the tidal volume. The normal tidal volume is about 500 mL. A great deal of stale air remains in the lungs during normal resting breathing. The volume of air that remains in the lungs at the end of a maximal expiration is the residual volume (about 1200 mL). The vital capacity is the maximum amount of air a person can exhale after filling the lungs to the maximum extent (about 4500 mL). Vital capacity is a useful measure of the functional capacity of the lungs.

Gas exchange takes place in the alveoli The respiratory system delivers oxygen to the alveoli, but if oxygen remained in the lungs, all the other body cells would soon die. The vital link between alveolus and body cell is the circulatory system. Pulmonary capillaries bring blood very close to the alveolar air.

Capillary in lung

(b) In the tissues. Oxygen is more concentrated in the blood than in the body cells; it diffuses out of the capillaries into the cells. Carbon dioxide is more concentrated in the cells, so it diffuses out of the cells and moves into the blood.

Cells in tissue PO = 40 mm Hg 2 PCO = 46 mm Hg 2

FIGURE 46-9 Gas exchange in the lungs and tissues Note the differences in partial pressures of oxygen and carbon dioxide before and after gases are exchanged in the tissues.

law of partial pressures, in a mixture of gases the total pressure of the mixture is the sum of the pressures of the individual gases. Each gas exerts, independently of the others, a partial pressure— the same pressure it would exert if it were present alone. At sea level, the barometric pressure (the pressure of Earth’s atmosphere) typically supports a column of mercury 760 mm high. Because oxygen makes up about 21% of the atmosphere, oxygen’s share of that pressure is 0.21 × 760 = 160 mm Hg. Thus, 160 mm Hg is the partial pressure of atmospheric O2, abbreviated PO2. In contrast, the partial pressure of atmospheric CO2 is 0.3 mm Hg, abbreviated PCO2. Fick’s law of diffusion explains that the amount of oxygen or carbon dioxide that diffuses across the membrane of an alveolus depends on the differences in partial pressure on the two sides of the membrane, and on the surface area of the membrane. The gas diffuses faster if the difference in pressure or the surface area increases.

Gas exchange takes place in the tissues The partial pressure of oxygen in arterial blood is about 100 mm Hg. The PO2 in the tissues is still lower, averaging about 40 mm Hg. Consequently, oxygen diffuses out of the capillaries and into the tissues. Not all the oxygen leaves the blood, however. The blood passes through the tissue capillaries too rapidly for equilibrium to be reached. As a result, the partial pressure of oxygen in venous blood returning to the lungs is about 40 mm Hg. Thus, exhaled air has had only part of its oxygen removed—a good thing for those in need of mouth-to-mouth resuscitation!

Respiratory pigments increase capacity for oxygen transport Respiratory pigments combine reversibly with oxygen and greatly increase the capacity of blood to transport it. Hemocyanins are copper-containing proteins dispersed in the hemolymph of many species of mollusks and arthropods. Without oxygen, these pigments are colorless. When oxygen combines with the copper, hemocyanins are blue. Hemoglobin and myoglobin are the most common respiratory pigments in animals. Hemoglobin is the pigment found in the blood of vertebrates. It is also present in many invertebrate species, including annelids, nematodes, mollusks, and arthropods. In some of these animals, the hemoglobin is dispersed in the plasma rather than confined to blood cells. Recall that myoglobin is a form of hemoglobin found in muscle fibers (see Chapter 40). In humans and other mammals, inhaled oxygen diffuses out of the alveoli and enters the pulmonary capillaries. Plasma in equilibrium with alveolar air can take up only 0.25 mL of oxygen per 100 mL. However, oxygen diffuses into red blood cells (RBCs) and combines with hemoglobin. The properties of hemoglobin permit whole blood to carry some 20 mL of oxygen per 100 mL. Hemoglobin transports almost 99% of the oxygen. The rest is dissolved in the plasma. The term hemoglobin is actually a general name for a group of related compounds, all of which consist of an iron–porphyrin, or heme, group bound to a protein known as a globin. The protein

portion of the molecule varies in size, amino acid composition, and physical properties among various species. The protein portion of hemoglobin is composed of four peptide chains, typically two a and two b chains, each attached to a heme (iron–porphyrin) ring (see Fig. 3-22a). An iron atom is bound in the center of each heme ring. Hemoglobin has the remarkable property of forming a weak chemical bond with oxygen. An oxygen molecule can attach to the iron atom in each heme. In the lung (or gill), oxygen diffuses into the RBCs and combines with hemoglobin (Hb) to form oxyhemoglobin (HbO2). When combined with oxygen, hemoglobin is bright red; without oxygen, it appears dark red, imparting a purplish color to venous blood. Because the chemical bond formed between the oxygen and the hemoglobin is weak, the reaction is readily reversible. As blood circulates through tissues where the oxygen concentration is low, the reaction proceeds to the left. Hemoglobin releases oxygen, which diffuses out of the blood and into the cells of the tissues. Hb + O2 Δ HbO2

The maximum amount of oxygen that hemoglobin can transport is its oxygen-carrying capacity. The actual amount of oxygen bound to hemoglobin is the oxygen content. The ratio of O2 content to oxygen-carrying capacity is the percent O2 saturation of the hemoglobin. The percent O2 saturation is highest in the pulmonary capillaries, where the concentration of oxygen is greatest. In the capillaries of the tissues, where there is less oxygen, oxyhemoglobin dissociates, releasing oxygen. There, the percent saturation of hemoglobin is correspondingly lower. The oxygen–hemoglobin dissociation curve shown in FIGURE 46-10a illustrates the relationship between partial pressure of oxygen and percent saturation of hemoglobin. As oxygen concentration increases, there is a progressive increase in the percentage of hemoglobin that is combined with oxygen. The ability of oxygen to combine with hemoglobin and be released from oxyhemoglobin is influenced by several factors in addition to percent O2 saturation. These factors include pH, carbon dioxide concentration, and temperature. Carbon dioxide produced in respiring tissue reacts with water in the plasma to form carbonic acid, H2CO3. For this reason, any increase in carbon dioxide concentration also increases the acidity (lowers the pH) of the blood. Oxyhemoglobin unloads oxygen more readily in an acidic environment than in an environment with normal pH. Displacement of the oxygen–hemoglobin dissociation curve by a change in pH is known as the Bohr effect (FIG. 46-10b). Lactic acid released from active muscles also lowers blood pH and has a similar effect on the oxygen–hemoglobin dissociation curve—more oxygen is unloaded and available for muscle contraction. Some carbon dioxide is transported by the hemoglobin molecule. Although carbon dioxide attaches to the hemoglobin molecule in a different way and at a different site than oxygen does, the attachment of a carbon dioxide molecule releases an oxygen molecule from the hemoglobin. This effect of carbon dioxide concentration on the oxygen–hemoglobin dissociation curve is important. In the capillaries of the lungs (or gills in fishes), carbon dioxide concentration is relatively low and oxygen concentration is high, so

KEY POINT

As oxygen concentration increases, the percentage of hemoglobin that combines with oxygen increases.

100

100

80 Oxygen-rich blood leaving the lungs

60 40

Oxygen-poor blood returning from tissues

20 0

0

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Partial pressure of oxygen (mm Hg)

Carbon dioxide is transported mainly as bicarbonate ions

(a) Normal oxygen–hemoglobin dissociation curve. Look at the relationship between the partial pressure of oxygen and the percent O2 saturation of hemoglobin. Oxygen binds to hemoglobin in the lungs and unloads from hemoglobin in the tissues.

Percent O2 saturation

Percent O2 saturation

oxygen combines with a very high percentage of hemoglobin. In the capillaries of the tissues, carbon dioxide concentration is high and oxygen concentration is low, so hemoglobin readily unloads oxygen. The oxygen and carbon dioxide bound to hemoglobin in RBCs do not contribute to the partial pressures that govern diffusion. Loading and unloading of gases onto hemoglobin depend on partial pressures of the plasma and interstitial fluid (which is determined by the gases dissolved in these fluids).

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(b) Effect of pH on the oxygen– hemoglobin curve (Bohr effect). The normal pH of human blood is 7.4. Find the location on the horizontal axis where the partial pressure of oxygen is 40, and follow the line up through the curves. Notice that the saturation of hemoglobin with oxygen differs among the three curves, even though the partial pressure of oxygen is the same. Oxygen loading increases at higher pH (7.6). At lower pH (7.2), hemoglobin unloads more oxygen.

Blood transports carbon dioxide in three forms. About 10% of carbon dioxide dissolves in plasma. Another 30% enters the RBCs and combines with hemoglobin. Because the bond between the hemoglobin and carbon dioxide is very weak, the reaction is readily reversFIGURE 46-10 Animated Oxygen–hemoglobin dissociation curves ible. Most carbon dioxide (about 60%) moves through plasma as bicarbonate ions (HCO3−). In plasma, carbon dioxide carbonic acid in the blood increases. When blood pH falls below 7, slowly combines with water to form carbonic acid. This reaction the central nervous system becomes depressed and the individual proceeds much more rapidly inside RBCs as a result of the action becomes disoriented. Untreated respiratory acidosis can cause of the enzyme carbonic anhydrase (FIG. 46-11). Carbonic acid dissociates, forming hydrogen ions and bicarbonate ions. coma and death.

CO2 Carbon dioxide

+

Carbonic anhydrase

H 2O Water

—¡ H2CO3 ¡ H+ + HCO3− Carbonic acid

Hydrogen ion

Bicarbonate ion

Most hydrogen ions released from carbonic acid combine with hemoglobin, which is a very effective buffer. Many of the bicarbonate ions diffuse into the plasma. The action of carbonic anhydrase in the RBCs maintains a diffusion gradient for carbon dioxide to move into and then out of RBCs. As the negatively charged bicarbonate ions move out of RBCs, chloride ions (Cl−) in the plasma diffuse into the RBCs to replace them, a process known as the chloride shift. In the alveolar capillaries, CO2 diffuses out of the plasma and into the alveoli. As the CO2 concentration decreases, the reaction sequence just described reverses. Any condition (such as emphysema) that interferes with the removal of carbon dioxide by the lungs can lead to respiratory acidosis. In this situation, carbon dioxide is produced more rapidly than it is excreted by the lungs. As a result, the concentration of

Breathing is regulated by respiratory centers in the brain Breathing is a rhythmic, involuntary process regulated by respiratory centers in the brain stem (see Fig. 46-6). Groups of neurons in the medulla regulate the basic rhythm of breathing. These neurons send a burst of impulses to the diaphragm and external intercostal muscles, which causes them to contract. After several seconds, these neurons become inactive, the muscles relax, and exhalation occurs. Respiratory centers in the pons help control the transition from inspiration to exhalation. These centers can stimulate or inhibit the respiratory centers in the medulla. The cycle of activity and inactivity repeats itself so that at rest you breathe about 14 times per minute. Overdose of certain medications, such as barbiturates, depresses the respiratory centers and may lead to respiratory failure. The basic rhythm of respiration changes in response to the body’s needs. When you are playing a fast game of tennis, you need

KEY POINT

The enzyme carbonic anhydrase (in red blood cells) converts carbon dioxide to carbonic acid; carbonic acid dissociates, producing hydrogen ions and bicarbonate ions. About 60% of carbon dioxide is transported in the plasma as bicarbonate ions. Plasma

Tissue cell CO2

Cl– Chloride shift

–

Cl–

Plasma

Red blood cell

H2O

Carbonic anhydrase H2CO3 Carbonic acid

Chloride shift Cl–

CO2 + H2O H2O CO2 H+

Hemoglobin

HCO3– + H+ Bicarbonate

H+ Hemoglobin

CO2

CO2 + H2O

Cl–

HCO3 + H+ Bicarbonate H2CO3 Carbonic acid

Tissue capillary wall

CO2

HCO3– Bicarbonate

CO2

CO2

Alveoli

CO2

Pulmonary capillary wall

CO2

HCO3– Bicarbonate

(a) In the tissues, carbon dioxide diffuses from cells into the plasma. Most of the carbon dioxide enters red blood cells, where the enzyme carbonic anhydrase rapidly converts it to carbonic acid. Carbonic acid dissociates, producing bicarbonate ions (HCO3–) and H+. As bicarbonate ions move out into the plasma, chloride ions replace them. Hemoglobin combines with H+, preventing a decrease in pH.

(b) In the lungs, carbon dioxide diffuses out of the plasma and into the alveoli. The reactions described in (a) are reversed. Bicarbonate ions diffuse from the plasma into the red blood cells. H+ released from hemoglobin combines with bicarbonate ions, producing carbonic acid. Carbon dioxide produced from the carbonic acid diffuses out of the blood and into the alveoli.

FIGURE 46-11 Carbon dioxide transport

more oxygen than when you are studying biology. During exercise the rate of aerobic cellular respiration increases, producing more carbon dioxide. Your body must dispose of this carbon dioxide through increased ventilation. Carbon dioxide concentration is the most important chemical stimulus for regulating the rate of respiration. Specialized chemoreceptors in the medulla and in the walls of the aorta and carotid arteries are sensitive to changes in arterial carbon dioxide concentration. When stimulated, they send impulses to the respiratory centers, which increase breathing rate. The chemoreceptors in the walls of the aorta, called aortic bodies, and those in the walls of the carotid arteries, the carotid bodies, are sensitive to changes in hydrogen ion concentration and oxygen concentration, as well as to carbon dioxide levels. Recall that an increase in carbon dioxide concentration increases hydrogen ions (from carbonic acid) and lowers blood pH. Even a slight decrease

in pH stimulates these chemoreceptors, leading to faster breathing. As the lungs remove carbon dioxide, the hydrogen ion concentration in the blood and other body fluids decreases, and homeostasis is re-established. Interestingly, oxygen concentration generally does not play an important role in regulating respiration. Only if the partial pressure of oxygen falls markedly do the chemoreceptors in the aorta and carotid arteries become stimulated to send messages to the respiratory centers. Shallow breathing, which occurs in anxiety and in many respiratory diseases, causes hypoxia, a deficiency of oxygen. Even rapid, shallow breathing results in hypoxia because we do not clear out the stale air in the airway and ventilate the lungs. Hypoxia causes drowsiness, mental fatigue, headache, and sometimes euphoria. The ability to think and make judgments is impaired, as is the ability to perform tasks requiring coordination.

Although breathing is involuntary, the action of the respiratory centers can be consciously influenced for a short time by stimulating or inhibiting them. For example, you can inhibit respiration by holding your breath. You cannot hold your breath indefinitely, however, because eventually you feel a strong urge to breathe. Even if you could ignore this urge, you would eventually pass out and resume breathing. People who have stopped breathing because of drowning, smoke inhalation, electric shock, or cardiac arrest can sometimes be sustained by mouth-to-mouth resuscitation until their own breathing reflexes return. Cardiopulmonary resuscitation (CPR) is a method for aiding victims who have suffered respiratory and cardiac arrest. CPR must be started immediately because irreversible brain damage occurs within about 4 minutes of oxygen deprivation. A number of organizations offer training in CPR to the general public.

Hyperventilation reduces carbon dioxide concentration Underwater swimmers and some Asian pearl divers undergo voluntary hyperventilation before going underwater. By making a series of deep inhalations and exhalations, they “blow off ” CO2, markedly reducing the CO2 content of the alveolar air and of the blood. This action allows them to remain underwater longer before the urge to breathe becomes irresistible. When hyperventilation continues for a long period, dizziness and sometimes unconsciousness may result. These responses occur because a certain concentration of carbon dioxide is needed in the blood to maintain normal blood pressure. (This mechanism operates by way of the vasoconstrictor center in the brain, which maintains the muscle tone of blood vessel walls.) Furthermore, if divers hold their breath too long, the low concentration of oxygen may result in unconsciousness and drowning.

High flying or deep diving can disrupt homeostasis The barometric pressure decreases at progressively higher altitudes. Because the concentration of oxygen in the air remains at 21%, the partial pressure of oxygen decreases along with the barometric pressure. When a person moves to a high altitude, his or her body adjusts over time by producing a greater number of RBCs. Getting sufficient oxygen from the air becomes an everincreasing problem as we ascend to progressively higher altitudes. All high-flying jets have airtight cabins pressurized to the equivalent of the barometric pressure at an altitude of about 2000 m (6600 ft). If a jet flying at 11,700 m (more than 38,000 ft) suddenly decompressed, the pilot would lose consciousness in about 30 seconds and become comatose in about 1 minute. In addition to the problems of hypoxia, a rapid decrease in barometric pressure causes decompression sickness (commonly known as the “bends” because those suffering from it bend over in pain). When the barometric pressure drops below the total pressure of all gases dissolved in the blood and other body fluids, the dissolved gases tend to come out of solution and form gas bubbles. A familiar example occurs each time you uncap a bottle of soda, reduc-

ing pressure in the bottle. Carbon dioxide is released from solution and bubbles out into the air. In the body, nitrogen has a low solubility in blood and tissues. When it comes out of solution, the bubbles produced may damage tissues and block capillaries, interfering with blood flow. The clinical effects of decompression sickness are pain, dizziness, paralysis, unconsciousness, and even death. Decompression sickness is more common in scuba diving than in high-altitude flying. As a diver descends, the surrounding pressure increases tremendously—1 atmosphere (the atmospheric pressure at sea level, which equals 760 mm Hg) for each 10 m (about 32 ft). To prevent lung collapse, a diver must be supplied with air under pressure, exposing the lungs to very high alveolar gas pressures. At sea level an adult human has about 1 L of nitrogen dissolved in the body, with about half in the fat and half in the body fluids. After a diver’s body has become saturated with nitrogen at a depth of 100 m (325 ft), the body fluids contain about 10 L of nitrogen. To prevent this nitrogen from rapidly bubbling out of solution and causing decompression sickness, the diver must rise to the surface gradually, with stops at certain levels on the way up. These pauses allow nitrogen to be expelled slowly through the lungs.

Some mammals are adapted for diving Some air-breathing mammals can spend long periods under water without coming up for air. Dolphins, whales, seals, and beavers have structural and physiological adaptations that allow them to dive for food or to elude their enemies. With their streamlined bodies and forelimbs modified as fins or flippers, diving mammals perform impressive aquatic feats. The Weddell seal can swim under the ice at a depth of 596 m (1955 ft) for more than an hour without coming up for air. The enormous elephant seal, which measures about 5 m (16 to 18 ft) in length and weighs 2 to 4 tons, can plunge even deeper (FIG. 46-12). A female elephant seal can dive to depths of more than 1500 m (almost 5000 ft) in less than 20 minutes and stay beneath the surface for more than an hour. Physiological adaptations, including ways to distribute and store oxygen, permit some mammals to dive deeply and remain underwater for long periods. Turtles and birds that dive depend on oxygen stored in their lungs. However, diving mammals do not take in and store extra air before a dive. In fact, seals exhale before they dive. With less air in their lungs, they are less buoyant. Their lungs collapse at about 50 to 70 m into their dive and then reinflate as they ascend, so their lungs do not function for most of the dive. These adaptations are thought to reduce the chance of decompression sickness because with less air in the lungs there is less nitrogen in the blood to dissolve during the dive. Seals have about twice the volume of blood, relative to their body weight, as nondiving mammals. Diving mammals also have high concentrations of myoglobin, which stores oxygen in muscles. These animals have up to ten times as much myoglobin as do terrestrial mammals. The very large spleen typical of many diving mammals stores oxygen-rich red blood cells. During a dive, the increased pressure squeezes the spleen, releasing the stored red blood cells into the circulation. Diving mammals markedly reduce the energy expended in deep diving (more than 200 m) by gliding. Filmed video sequences

in which the victim was submerged for as long as 45 minutes in very cold water before being rescued and resuscitated. Many of these survivors showed no brain damage. The shock of icy water slows heart rate, increases blood pressure, and shunts blood to internal organs of the body that most need oxygen (blood flow in arms and legs decreases). Metabolic rate decreases, so less oxygen is required.

of diving seals and whales show that they glide most of the way down. Gliding is possible because the animal’s buoyancy decreases as the lungs gradually collapse and the amount of air in the lungs is reduced. Gravity then pulls the animal downward. When a mammal dives to its limit, physiological mechanisms known collectively as the diving reflex are activated. Metabolic rate decreases by about 20%, which conserves oxygen. Breathing stops, and bradycardia (slowing of the heart rate) occurs. The heart rate may decrease to one-tenth the normal rate, reducing the body’s consumption of oxygen and energy. Blood is redistributed so that the heart and brain receive the greatest share; skin, muscles, digestive organs, and other internal organs can survive with less oxygen, and less blood is directed to these organs while an animal is submerged. The diving reflex is present to some extent in humans, where it may act as a protective mechanism during birth, when an infant may be deprived of oxygen for several minutes. Cases of near drownings, especially of young children, have been documented

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What is the sequence of inhaled airflow through the respiratory structures in a mammal? What happens in the alveoli? Why does alveolar air differ in composition from atmospheric air? Explain. How does carbon dioxide concentration affect the oxygen–hemoglobin dissociation curve? What physiological mechanisms bring about an increase in rate and depth of breathing during exercise? Why is such an increase necessary?

46.4 BREATHING POLLUTED AIR ■ ■ LEARNING OBJECTIVE 7

Describe the defense mechanisms that protect the lungs and describe the effects of polluted air and cigarette smoking on the respiratory system.

Bruce Watkins/Animals Animals

David Nunuk/Science Photo Library/Photo Researchers, Inc.

Several defense mechanisms protect the delicate lungs from the harmful substances we breathe (FIG. 46-13). Hairs in the nostrils, the ciliated mucous lining in the nose and pharynx, and the cilia– mucus elevator of the trachea and bronchi trap foreign particles in inspired air. Researchers recently reported that cilia of epithelial cells lining the human airway function as chemoreceptors that sense bitter taste. Bitter compounds increase the concentration of Ca2+, which, in turn, increase the rate of ciliary beating. As a result the harmful compound is more rapidly removed.

FIGURE 46-12 Deep diver Elephant seals, like this northern elephant seal (Mirounga angustirostris), may be the deepest divers on Earth. Researchers have recorded their dives at depths of more than 1500 m (almost 5000 ft).

FIGURE 46-13 Urban air pollution Industry continues to spew tons of pollutants into the atmosphere. Despite modern technology, air pollution still poses a significant health risk.

In qu i ri n g A b o u t THE EFFECTS OF SMOKING What is the single most preventable cause of disease, disability, and death in the United States? According to the Centers for Disease Control, it is tobacco use. Cigarette smoking kills more people than alcohol, car accidents, suicide, AIDS, homicide, and illegal drugs combined. More than 443,000 people die prematurely each year from smoking or exposure to secondhand smoke. An additional 8.6 million have a serious illness caused by smoking.1 Tobacco smoke is by far the most important risk factor for lung cancer, the most common lethal cancer worldwide. It is also associated with many other types of cancer and accounts for at least 30% of all cancer deaths. Tobacco smoke is also an important risk factor for cardiovascular disease, chronic obstructive pulmonary disease, and many other diseases. Despite these grim facts, approximately 20% of both high school students and adults in the United States continue to smoke cigarettes. (This amounts to more than 43 million U.S. adults and 71 million teens.) Secondhand tobacco smoke is a known human carcinogen. It causes the deaths of an estimated 49,000 individuals each year in the United States alone. Millions of children and adults are exposed to secondhand smoke in their homes and workplaces. This exposure causes disease and premature death in children and adults who do not smoke. Babies and children exposed to secondhand smoke are at increased risk for sudden infant death syndrome (SIDS), acute respiratory infections, pneumonia, bronchitis, ear infections, and asthma. In adults, secondhand smoke causes cardiovascular disease and lung cancer. About 60% of nonsmokers in the United States exhibit biological evidence of exposure to secondhand smoke.

1

Statistics quoted are from the Centers for Disease Control and the American Cancer Society.

Nicotine is highly addictive. Like morphine, amphetamines, and cocaine, nicotine increases dopamine concentration in the limbic system. This area helps integrate emotion, and dopamine may facilitate learning an association between the pleasurable effects of the drug and other stimuli, such as the smell of smoke. Nicotine, like cocaine and other abused drugs, causes long-lasting changes in the brain. Cell signaling and other mechanisms underlying drug addiction are very similar to those of learning and memory. An estimated 70% of smokers (more than 33 million) want to quit, but only 2.5% per year succeed in quitting permanently. Nicotine replacement with gum, patches, nasal spray, inhalers, or lozenges has been shown to be effective as an aid to smoking cessation, especially when used with behavioral therapy. Certain antidepressants are used to reduce the craving for nicotine. A newer prescription medication (Chantix) works by blocking nicotine receptors in the brain. It reduces symptoms of nicotine withdrawal and also decreases the pleasurable effects of smoking. Here are some facts about smoking: • Smoking shortens the life of an adult male smoker by an average of 13.2 years and shortens the life of an adult female smoker by an average of 14.5 years. • If you smoke more than one pack per day, you are about 20 times as likely as a nonsmoker to develop lung cancer. According to the American Cancer Society, cigarette smoking causes almost nine out of ten lung cancer deaths. • If you smoke, you double your chances of dying from cardiovascular disease. • If you smoke, you are 20 times as likely as a nonsmoker to develop chronic bronchitis and emphysema. About 90% of all deaths from COPD are caused by cigarette smoking.

One of the body’s most rapid defense responses to breathing dirty air is bronchial constriction. In this process, the bronchial tubes narrow, increasing the chance that inhaled particles will land on the sticky mucous lining. Unfortunately, bronchial constriction narrows the airway. Less air can reach the lungs, decreasing the amount of oxygen available to body cells. Fifteen puffs on a cigarette during a 5-minute period increases airway resistance as much as threefold, and this added resistance to breathing lasts more than 30 minutes. Chain-smokers and those who breathe heavily polluted air are in a state of chronic bronchial constriction.

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•

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If you smoke, you have about 5% less oxygen circulating in your blood (because carbon monoxide binds to hemoglobin) than does a nonsmoker. If you smoke when you are pregnant, your baby will weigh about 6 ounces less at birth, and there is a higher risk of miscarriage, premature birth (early delivery), stillbirth, and sudden infant death syndrome. Infants whose parents smoke have double the risk of contracting pneumonia or bronchitis in their first year of life. There is no risk-free level of exposure to secondhand smoke. Nonsmokers who are exposed to secondhand smoke increase their risk of developing heart disease by 25% to 30% and lung cancer by 20% to 30%. When smokers quit smoking, their risk of dying from chronic obstructive pulmonary disease, cardiovascular disease, or cancer gradually decreases. (Precise changes in risk depend on the number of years the person smoked, the number of cigarettes smoked per day, the age of starting to smoke, and the number of years since quitting.) Almost every American who takes up smoking is a teenager—more than 3500 teens smoke their first cigarette every day. About one-third of these teens will die prematurely from a smoking-related disease. The younger you are when you begin to smoke, the more likely you are to be an adult smoker. The direct medical costs of tobacco use amount to more than $96 billion annually. Smokeless tobacco products (snuff, chewing tobacco) are not a safe alternative to tobacco smoking. All forms of these products have chemicals that cause cancer.

Neither the smallest bronchioles nor the alveoli are equipped with mucus or ciliated cells. Foreign particles that get through other respiratory defenses and find their way into the alveoli may be engulfed by macrophages. The macrophages may then accumulate in the lymph tissue of the lungs. Lung tissues of cigarette smokers and those who work in dirty industries burning fossil fuel contain large blackened areas where carbon particles have been deposited (FIG. 46-14). Continued insult to the respiratory system results in disease. Chronic bronchitis, pulmonary emphysema, and lung cancer have

Martin Rotker

Martin Rotker

(a) Lungs and major bronchi of a nonsmoker.

physema, a disease also most common in cigarette smokers. In this disorder, alveoli lose their elasticity, and walls between adjacent alveoli are destroyed. The surface area of the lung is so reduced that gas exchange is seriously impaired. Air is not expelled effectively, and stale air accumulates in the lungs. The emphysema victim struggles for every breath, and still the body does not get enough oxygen. To compensate, the right ventricle of the heart pumps harder and becomes enlarged. Emphysema patients frequently die of heart failure. Most patients with chronic obstructive pulmonary disease (COPD), a condition characterized by obstructed airflow, have both chronic bronchitis and emphysema. Asthma also contributes to COPD. People with asthma respond to inhaled stimuli with exaggerated bronchial constriction, and the airway is typically inflamed. Cigarette smoking is also the main cause of lung cancer. More than 69 of the 4800 chemical compounds in tobacco smoke cause cancer in animals, including humans. These carcinogenic substances irritate the cells lining the respiratory passages and alter their metabolic balance. Normal cells are transformed into cancer cells, which may multiply rapidly and invade surrounding tissues.

(b) Lungs and heart of a cigarette smoker. The dark spots visible in the lung tissue are particles of carbon, tar, and other substances that passed through the respiratory defenses and lodged in the lungs.

FIGURE 46-14 Effects of cigarette smoking

Review been linked to cigarette smoking and breathing polluted air. More than 75% of patients with chronic bronchitis have a history of heavy cigarette smoking (see Inquiring About: The Effects of Smoking). People with chronic bronchitis often develop pulmonary em-

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What mechanisms does the human respiratory system have for getting rid of inhaled particles? What happens when so much dirty air is inhaled that these mechanisms cannot function effectively?

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

46.1 (page 994) 1 Compare the advantages and disadvantages of air and water as mediums for gas exchange, and describe adaptations for gas exchange in air. ■ Air contains a higher concentration of molecular oxygen than does water, and oxygen diffuses more rapidly through air than through water. Air is less dense and less viscous than water, so less energy is needed to move air over a gas exchange surface. ■ Terrestrial animals have adaptations that protect their respiratory surfaces from drying.

46.2 (page 994) 2 Describe the following adaptations for gas exchange: body surface, tracheal tubes, gills, and lungs. Small aquatic animals exchange gases by diffusion; they require no specialized respiratory structures. Some invertebrates, including most annelids, and a few vertebrates (many amphibians) exchange gases across the body surface. ■ In insects and some other arthropods, air enters a network of tracheal tubes, or tracheae, through openings, called spiracles, along the body surface. Tracheal tubes branch and extend to all regions of the body. ■ Gills are moist, thin projections of the body surface found mainly in aquatic animals. In chordates, gills are usually internal, located along the edges of the gill slits. In bony fishes an operculum pro-

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tects the gills. A countercurrent exchange system maximizes diffusion of oxygen into the blood and diffusion of carbon dioxide out of blood. Animals carry on ventilation, the process of actively moving air or water over respiratory surfaces. Terrestrial vertebrates have lungs and some means of ventilating them. In birds, the lungs have extensions, called air sacs, that act as bellows, drawing air into the system. Two cycles of inhalation and exhalation support a one-way flow of air through the lungs. Air from the outside flows into the posterior air sacs, to the lung, through the anterior air sacs, and then out of the body. Gas exchange takes place through the walls of the parabronchi, small, open tubes in the lungs. A crosscurrent arrangement, in which blood flow is at right angles to the parabronchi, increases the amount of oxygen that enters the blood.

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Learn more about adaptations for gas exchange, including gills in bony fishes, vertebrate lungs, and the bird respiratory system, by clicking on the figures in CengageNOW.

46.3 (page 998) 3 Trace the passage of oxygen through the human respiratory system from nostrils to alveoli. ■ The human respiratory system includes the lungs and a system of airways. Each breath of air passes in sequence through the nostrils, nasal cavities, pharynx, larynx, trachea, bronchi, bronchioles,

and alveoli. Each lung occupies a pleural cavity and is covered with a pleural membrane. Bronchiole

Alveolus

Capillaries Alveolus

Alveolus

Learn more about the human respiratory system by clicking on the figures in CengageNOW.

4 Summarize the mechanics of breathing in humans, and describe gas exchange in the lungs and tissues. During breathing, the diaphragm contracts, expanding the chest cavity. The membranous walls of the lungs move outward along with the chest walls, lowering pressure within the lungs. Air from outside the body rushes in through the air passageways and fills the lungs until the pressure equals atmospheric pressure. ■ Tidal volume is the amount of air moved into and out of the lungs with each normal breath. The volume of air that remains in the lungs at the end of a maximal expiration is the residual volume. Vital capacity is the maximum volume of air that can be exhaled after the lungs fill to the maximum extent. ■ Oxygen and carbon dioxide are exchanged between alveoli and blood by diffusion. The pressure of a particular gas determines its direction and rate of diffusion. ■ Dalton’s law of partial pressures explains that in a mixture of gases, the total pressure of the mixture is the sum of the pressures of the individual gases. Thus, each gas in a mixture of gases exerts a partial pressure, the same pressure it would exert if it were present alone. The partial pressure of atmospheric oxygen, PO2, is 160 mm Hg at sea level. According to Fick’s law of diffusion, the greater the difference in ■ pressure on the two sides of a membrane and the larger the surface area, the faster the gas diffuses across the membrane. ■

See the breathing mechanisms in action by clicking on the figures in CengageNOW.

5 Summarize the mechanisms by which oxygen and carbon dioxide are transported in the blood and identify factors that determine the oxygen–hemoglobin dissociation curve.

Hemoglobin is the respiratory pigment in the blood of vertebrates. Almost 99% of the oxygen in human blood is transported as oxyhemoglobin (HbO2). ■ The maximum amount of oxygen that hemoglobin can transport is the oxygen-carrying capacity. The actual amount of oxygen bound to hemoglobin is the oxygen content. The percent O2 saturation, the ratio of oxygen content to oxygen-carrying capacity, is highest in pulmonary capillaries, where oxygen concentration is greatest. ■ The oxygen–hemoglobin dissociation curve shows that as oxygen concentration increases, there is a progressive increase in the amount of hemoglobin that combines with oxygen. The curve is affected by pH, temperature, and CO2 concentration. ■ Oxyhemoglobin dissociates more readily as carbon dioxide concentration increases because carbon dioxide combines with water and produces carbonic acid, which lowers the pH. Displacement of the oxygen–hemoglobin dissociation curve by a change in pH is called the Bohr effect. ■ About 60% of the carbon dioxide in the blood is transported as bicarbonate ions. About 30% combines with hemoglobin, and another 10% is dissolved in plasma. ■ Carbonic anhydrase catalyzes the production of carbonic acid from carbon dioxide and water. Carbonic acid dissociates, producing bicarbonate ions (HCO3−) and hydrogen ions (H+). ■ Hemoglobin combines with H+, which buffers the blood. Many bicarbonate ions diffuse from red blood cells into the plasma; they are replaced by Cl−. This exchange is known as the chloride shift. 6 Describe the regulation of breathing in humans and summarize the physiological effects of hyperventilation and of sudden decompression when a diver surfaces too quickly from deep water. ■ Respiratory centers in the medulla and pons regulate respiration. These centers are stimulated by chemoreceptors sensitive to an increase in carbon dioxide concentration. They also respond to an increase in hydrogen ions and to very low oxygen concentration. ■ Hyperventilation reduces the concentration of carbon dioxide in the alveolar air and in the blood. ■ As altitude increases, barometric pressure falls, and less oxygen enters the blood. This situation can lead to hypoxia, or oxygen deficiency, which in turn can lead to loss of consciousness and death. Rapid decrease in barometric pressure can also cause decompression sickness, especially common among divers who ascend too rapidly from a dive. ■ Diving mammals have high concentrations of myoglobin, a pigment that stores oxygen, in their muscles. The diving reflex, a group of physiological mechanisms, including a decrease in metabolic rate, is activated when a mammal dives to its limit. ■

46.4 (page 1007) 7 Describe the defense mechanisms that protect the lungs and describe the effects of polluted air and cigarette smoking on the respiratory system. ■ The ciliated mucous lining of the airway traps inhaled particles. Inhaling polluted air results in bronchial constriction, increased mucus secretion, damage to ciliated cells, and coughing. Breathing polluted air or inhaling cigarette smoke can cause many diseases, including chronic bronchitis, pulmonary emphysema, and lung cancer.

T E S T YO U R U N D E R S TA N D I N G 1. Which of the following is a benefit of gas exchange in air compared with gas exchange in water? (a) air has a higher concentration of molecular oxygen (b) oxygen diffuses more slowly in air (c) no energy is required for ventilation (d) moist respiratory surface is not needed (e) air is denser than water 2. Which of the following are accurately matched? (a) bony fish— operculum (b) insect—alveoli (c) bird—spiracles (d) aquatic mammal—gill filaments (e) shark—dermal gills 3. Tracheal tubes (tracheae) (a) are linked to the circulatory system (b) are highly vascular (c) branch and extend to all the cells (d) are characteristic of many vertebrates (e) a, b, and c 4. The most efficient vertebrate respiratory system is that of (a) amphibians (b) birds (c) reptiles (d) mammals (e) humans

9. The concentration of which substance is most important in regulating the rate of respiration? (a) chloride ions (b) oxygen (c) bicarbonate ions (d) nitrogen (e) carbon dioxide 10. When a diver ascends too rapidly, (a) bronchial constriction occurs (b) a diving reflex is activated (c) nitrogen rapidly bubbles out of solution in the body fluids (d) nitrogen hypoxia occurs (e) carbon dioxide bubbles damage the alveoli 11. Which of the following is not true of the diving reflex? (a) breathing stops (b) the heart slows (c) less blood is distributed to the muscles (d) metabolic rate increases by about 20% (e) energy consumption decreases 12. Label the figure. Use Figure 46-6 to check your answers.

5. In a bird, the correct sequence for a breath of air is 1. anterior air sacs 2. posterior air sacs 3. parabronchi 4. alveoli (a) 1, 2, 4 (b) 2, 1, 3 (c) 2, 3, 1 (d) 2, 4, 1 (e) 1, 3, 2 6. Which sequence most accurately describes the sequence of airflow in the human respiratory system? 1. pharynx 2. bronchus 3. trachea 4. larynx 5. alveolus 6. bronchiole (a) 4, 1, 3, 2, 5, 6 (b) 1, 4, 3, 2, 5, 6 (c) 4, 1, 3, 2, 6, 5 (d) 1, 4, 3, 2, 6, 5 (e) 1, 3, 4, 2, 6, 5 7. The amount of air moved in and out of the lungs with each normal resting breath is the (a) vital capacity (b) residual volume (c) vital volume (d) partial pressure (e) tidal volume 8. According to Fick’s law of diffusion, a gas will diffuse faster if the (a) surface is reduced (b) difference in pressure is decreased (c) difference in pressure is increased (d) pH is lowered (e) a and c

CRITICAL THINKING 1. What problems would be faced by a terrestrial animal having gills instead of lungs? 2. EVOLUTION LINK. Under what conditions might it be advantageous for a fish to have lungs as well as gills? What function do the “lungs” of modern fishes serve? 3. EVOLUTION LINK. Aquatic mammals such as whales and dolphins use lungs rather than gills for gas exchange. Propose a hypothesis to explain this. 4. EVOLUTION LINK. What are the advantages of having millions of alveoli rather than a pair of simple, balloonlike lungs? 5. ANALYZING DATA. Look at Figure 46-10b. How would strenuous muscle activity affect the oxygen–hemoglobin disso-

ciation curve? (Hint: How does muscular activity affect pH? See Chapter 40 if you need to review this concept.) What would be the advantage of the shift in the curve? 6. SCIENCE, TECHNOLOGY, AND SOCIETY. How would you solve the tobacco smoking problem, which is extremely expensive to our society in economic and health costs, as well as in human suffering? Additional questions are available in CengageNOW at www.cengage.com/ login.

Processing Food and Nutrition

Jonathan Bird/Peter Arnold, Inc.

47

Atlantic wolffish (Anarhichas lupus) eating a sea urchin. Also known as the seawolf and as the Atlantic catfish, this fish lives on the ocean floor where it hunts and eats hard-shelled prey. The Atlantic wolffish can weigh up to 40 pounds. Its population has decreased drastically as a result of overfishing, unintentional harm by fishermen (bycatch), and habitat damage. Photographed in Maine.

KEY CONCEPTS

S

47.1 Many animal adaptations are associated with mode

some insects capture other animals. Although their choices of food and

of nutrition; food processing includes ingestion, digestion, absorption, and elimination.

feeding mechanisms are diverse, all animals are heterotrophs, organisms

47.2 Various parts of the vertebrate digestive system are

ecules manufactured by other organisms. What they eat, how they obtain

specialized to perform specific functions; accessory glands (liver, pancreas, and salivary glands) secrete fluids and enzymes that are important in digestion.

ponges and baleen whales feed on food particles suspended in the water. Giraffes eat leaves, and antelope graze on grasses. Frogs, lions, and

that must obtain their energy and nourishment from the organic molfood, and how they process and use food are the focus of this chapter. Nutrients are substances in food that are used as energy sources

47.3 Most animals require the same basic nutrients:

to power the systems of the body, as ingredients to make compounds

carbohydrates, lipids, proteins, vitamins, and minerals.

needed for metabolic processes, and as building blocks in the growth and

47.4 Basal metabolic rate is the body’s cost of metabolic

repair of tissues. Obtaining nutrients is of such vital importance that both

living; energy metabolism is regulated by complex signaling pathways. Both overnutrition and undernutrition can result in malnutrition.

individual organisms and ecosystems are structured around the central theme of nutrition, the process of taking in and using food. An animal’s body plan and its lifestyle are adapted to its particular mode of obtaining food. For example, the seawolf lives on the ocean floor at depths of up to 120 m (400 ft). It hunts clams, crabs, sea stars, and sea urchins (see photograph). Both its upper and lower jaw have four to six fanglike teeth that stab and tear its prey. Behind these fangs, there are

three rows of teeth that crush the hard shells of its prey. It also has serrated teeth in its throat. Nutrition has been an important force in human evolution. Through natural selection, the human diet became more varied than the diets of other primates. Humans also became very efficient at obtaining food. Their higher-quality diet supported the evolution of a larger, more complex brain. With only slight variations, all animals require the same basic nutrients: carbohydrates, lipids, proteins, vitamins, and minerals. Carbohydrates, lipids, and proteins are used as energy sources. Eating too much of any of these nutrients can result in weight gain, whereas eating too few nutrients or an unbalanced diet can result in malnutrition and death. Malnutrition, or poor nutritional status, results from dietary intake that is either below or above required needs. In human populations, both undernutrition (particularly protein deficiency) and obesity (which results from overnutrition) are serious health problems. Food processing and nutrition are active areas of research.

47.1 NUTRITIONAL STYLES AND ADAPTATIONS ■ ■ LEARNING OBJECTIVE 1

Describe food processing, including ingestion, digestion, absorption, and egestion or elimination, and compare the digestive system of a cnidarian (such as Hydra) with that of an earthworm and vertebrate.

Feeding is the selection, acquisition, and ingestion of food. Most animals have a specialized digestive system that processes the food they eat. Ingestion is the process of taking food into the digestive cavity. In many animals, including vertebrates, ingestion includes taking food into the mouth and swallowing it. The process of breaking down food is called digestion. Because animals eat the macromolecules made by and for other organisms, they must break down these molecules and synthesize new macromolecules for their own needs. For example, we humans cannot incorporate the proteins and other organic compounds from a hamburger directly into our own cells. We must mechanically digest the meat and then chemically digest it by enzymatic hydrolysis (see Chapter 3). During digestion, complex organic compounds are degraded into smaller molecular components. For example, proteins are broken down into their component amino acids. These amino acids will later be used to synthesize proteins tailored for the needs of that individual at that particular time. Amino acids and other nutrients pass through the lining of the digestive tract and into the blood by absorption. Then the circulatory system transports the nutrients to all the cells of the body. Cells use nutrients to synthesize proteins and other organic compounds, or as fuel for cellular respiration. Food that is not digested and absorbed is discharged from the body, a process called egestion or elimination.

Animals are adapted to their mode of nutrition We classify animals as herbivores, carnivores, or omnivores on the basis of the type of food they typically eat (FIG. 47-1). Many adaptations have evolved for each type of feeding. For example, the beaks of birds and the teeth of many vertebrates are specialized for cutting, tearing, crushing, or chewing food. Animals that feed directly on producers are herbivores, or primary consumers. Most of what an herbivore eats is not efficiently digested and is eliminated from the body, almost unchanged, as waste. For this reason herbivores must eat large quantities of food to obtain the nourishment they need. Many herbivores— grasshoppers, locusts, elephants, and cattle, for example—spend a major part of their lives eating. Animals cannot digest the cellulose of plant cell walls, and many adaptations have evolved for extracting nutrients from the plant material they eat. Many herbivores, including termites, cows, and horses, have a symbiotic relationship with bacteria, fungi, and other microorganisms that inhabit their digestive tracts. Ruminants (cattle, sheep, deer, giraffes) are hooved animals with a stomach divided into four chambers. Symbiotic bacteria and protists living in the first two chambers digest cellulose, splitting some of the molecule into sugars, which are then used by the host and the bacteria themselves. The bacteria produce fatty acids during their metabolism, some of which are absorbed by the animal and serve as an important energy source. Solid bits of food clump together to form a cud. The cud is regurgitated into the animal’s mouth, where it is mixed with saliva and chewed again. When the cud is reswallowed, the partly digested food is further broken down. Partially digested food passes into the third chamber, where water and minerals are absorbed into the blood. Digestion is completed in the fourth chamber, which corresponds to the stomach of humans and other animals with a single stomach. Herbivores are sometimes eaten by flesh-eating carnivores, which may also eat one another. Many carnivores (secondary and higher-level consumers in ecosystems) are predators, adapted for capturing and killing prey. Some carnivores seize their victims and swallow them alive and whole (see Fig. 47-1d). Others paralyze, crush, or shred their prey before ingesting it. Adaptations for this type of feeding include tentacles, claws, fangs, poison glands, and teeth. Carnivorous mammals have well-developed canine teeth for stabbing their prey during combat. Spotted hyenas, which are formidable predators as well as scavengers, have massive jaws that allow them to consume an entire elephant carcass, including bones and hide. Few nutrients are wasted. Meat is more easily digested than plant food, and carnivore digestive tracts are shorter than those of herbivores. Omnivores, such as pigs, humans, most bears, and some fishes, consume both plants and animals. Earthworms ingest large amounts of soil containing both animal and plant material. The blue whale, the largest animal, filters out tiny algae and animals from the water as it swims. Omnivores often have adaptations that help them distinguish among a wide range of smells and tastes and thereby select a variety of foods.

Darwin Dale / Photo Researchers, Inc.

Tom McHugh / Photo Researchers Inc.

(b) An herbivorous bear. The giant panda’s (Ailuropoda melanoleuca) large, flat teeth and well-developed jaws and jaw muscles are adaptations for grinding high-fiber plant food. Most bears are omnivores.

Frans Lanting /Minden Pictures

Carmela Lesczynski / Animals Animals / Earth Scenes

(a) Unusual adaptation of an acorn-eating insect. The impressively long “snout” of the herbivorous acorn weevil (Curculio) is adapted both for feeding and for making a hole in the acorn through which it deposits an egg. When it has hatched, the larva feeds on the contents of the acorn seed.

(c) A carnivorous fish. The mouth (on left ) of the longnose butterfly fish (Forcipiger longirostris) is adapted for extracting small worms and crustaceans from tight spots in coral reefs.

(d) Snakes are carnivorous. This snake (Dromicus) is strangling a lava lizard (Tropidurus ). A snake can swallow very large prey whole because of the structure of the jaws. The very flexible lower jaw expands during swallowing.

FIGURE 47-1 Adaptations for obtaining and processing food

Animals can also be classified according to the mechanisms they use to feed. Many omnivores are filter feeders that remove suspended food particles from the water they inhabit. For example, clams and oysters trap suspended food particles in mucus secreted by their gills. Some echinoderms trap food particles along their tentacles, which are coated with mucus. Baleen whales use rows of

hard plates (baleen) suspended from the roof of the mouth to filter small crustaceans. Some animals feed on fluids by piercing and sucking. Mosquitoes have highly adapted structures for piercing skin and sucking blood. Bats that feed on nectar have a long tongue and reduced dentition (number of teeth). Birds that feed on pollen and nectar have long bills

▲

FIGURE 47-2 Animated Simple invertebrate digestive systems

Wastes

(a) Hydras and (b) flatworms (planarians) have a gastrovascular cavity, a digestive tract with a single opening that serves as both mouth and anus. Food

Tentacle Mouth

Food

Enzymes secreted by gastrodermis (inner layer)

Gastrovascular cavity

Food absorbed Gastrodermis

(a) Digestive system of a hydra.

Gastrovascular cavity

Epidermis

Food particles Pharynx Food absorbed

Mouth

Enzymes Lining of the intestine

Wastes Food

(b) Digestive system of a flatworm.

and tongues. The shape, size, and curve of the beak may be specialized for feeding on or capturing a particular type of food (see Fig. 1-11).

Some invertebrates have a digestive cavity with a single opening Sponges, very simple invertebrates, obtain food by filtering microscopic organisms from the surrounding water. Individual cells phagocytose the food particles, and digestion is intracellular within food vacuoles. Wastes are egested into the water that continuously circulates through the sponge body. Most animals have a digestive cavity. Digestion within a cavity is more efficient than intracellular digestion because digestive enzymes can be released into one confined space. Less surface area is required. Cnidarians (such as hydras and jellyfish) and flatworms

have a gastrovascular cavity, a central digestive cavity with a single opening. Cnidarians capture small aquatic animals with the help of their stinging cells and tentacles (FIG. 47-2a). The mouth opens into the gastrovascular cavity. Cells lining this digestive cavity secrete enzymes that break down proteins. Digestion continues intracellularly within food vacuoles, and digested nutrients diffuse into other cells. Body contractions promote egestion of undigested food particles through the mouth. Free-living flatworms begin to digest their prey even before ingesting it. They extend the pharynx out through their mouth and secrete digestive enzymes onto their prey (FIG. 47-2b). After it is ingested, the food enters the branched gastrovascular cavity, where enzymes continue to digest it. Cells lining the gastrovascular cavity phagocytose partly digested food fragments, and digestion is completed within food vacuoles. As in cnidarians, the flatworm digestive cavity has only one opening, so undigested wastes are egested through the mouth.

Most animal digestive systems have two openings Most invertebrates, and all vertebrates, have a tube-withina-tube body plan. The body wall forms the outer tube. The inner tube is a digestive tract with two openings, sometimes referred to as a complete digestive system (FIG. 47-3). Food enters through the mouth, and undigested food is eliminated through the anus. The mixing and propulsive movements of the digestive tract are referred to as motility. The propulsive activity characteristic of most regions of the digestive tract is peristalsis, waves of muscular contraction that push the food in one direction. More food can be taken in while

Crop

Gizzard

Esophagus Pharynx Mouth Food

Intestine Anus Wastes

FIGURE 47-3 Digestive tract with two openings The earthworm, like most animals, has a complete digestive tract extending from mouth to anus. Various regions of the digestive tract are specialized to perform different food-processing functions.

previously eaten food is being digested and absorbed farther down the digestive tract. In a digestive tract with two openings, various regions of the tube are adapted to perform specific functions.

Review ■ ■ ■

How have carnivores adapted to their mode of nutrition? How does food processing differ in earthworms and flatworms? What are the advantages of a digestive tract with two openings?

47.2 THE VERTEBRATE DIGESTIVE SYSTEM ■ ■ LEARNING OBJECTIVES 2

3

Trace the pathway traveled by an ingested meal through the human digestive system and describe the structure and function of each organ involved. Trace the step-by-step digestion of carbohydrate, protein, and lipid.

4 Describe the structural adaptations that increase the surface area of the 5

digestive tract. Compare lipid absorption with absorption of other nutrients.

Various regions of the vertebrate digestive tract are specialized to perform specific functions (FIG. 47-4). Food passes in sequence through the following specialized regions: mouth ¡ pharynx (throat) ¡ esophagus ¡ stomach ¡ small intestine ¡ large intestine ¡ anus

The liver, pancreas, and the salivary glands (in terrestrial vertebrates) are accessory glands that secrete digestive juices into the digestive tract. The wall of the digestive tract has four layers. Although various regions differ somewhat in structure, the layers are basically similar throughout the digestive tract (FIG. 47-5). The mucosa, which consists of a layer of epithelial tissue and underlying connective tissue, lines the lumen (inner space) of the digestive tract. In the stomach and intestine, the mucosa is greatly folded to increase the

Parotid salivary gland Pharynx Esophagus

Liver Gallbladder Ascending colon

Sublingual salivary gland Submandibular salivary gland

Stomach Duodenum Pancreas Transverse colon Jejunum

lleum Cecum

Descending colon Sigmoid colon

Vermiform appendix

FIGURE 47-4 Animated Human digestive system The human digestive tract is a long, coiled tube extending from mouth to anus. The small intestine consists of the duodenum, jejunum, and ileum. The large intestine includes the cecum, colon, rectum, and anus. Locate the three types of accessory glands: the liver, pancreas, and salivary glands.

Rectum Anus

VISCERAL PERITONEUM

KEY POINT

An animal’s teeth are adapted for its mode of nutrition.

Lymph nodule Villi

Incisors

Blood vessels SUBMUCOSA Nerve fibers

MUCOSA

MUSCLE LAYER

Inner circular fibers Outer longitudinal fibers

Canines Premolars Molars

(a) Carnivore. The skull of a coyote shows the pointed incisors and canines, adaptations for ripping flesh.

Canine

FIGURE 47-5 Wall of the digestive tract From inside out, the layers of the wall are the mucosa, submucosa, muscle layer, and visceral peritoneum.

secreting and absorbing surface. Surrounding the mucosa is the submucosa, a connective tissue layer rich in blood vessels, lymphatic vessels, and nerves. A muscle layer, consisting of two sublayers of smooth muscle, surrounds the submucosa. In the inner sublayer, the muscle fibers are arranged circularly around the digestive tube. In the outer sublayer, the muscle fibers are arranged longitudinally. Below the level of the diaphragm, the outer connective tissue coat of the digestive tract is called the visceral peritoneum. By various folds it is connected to the parietal peritoneum, a sheet of connective tissue that lines the walls of the abdominal and pelvic cavities. The visceral and parietal peritonea enclose part of the coelom, the peritoneal cavity. Inflammation of the peritoneum, called peritonitis, can be very serious because infection can spread along the peritoneum to most of the abdominal organs.

Food processing begins in the mouth Imagine that you have just taken a big bite of a hamburger in a bun. The mouth is specialized for ingestion and for beginning the digestive process. Mechanical digestion begins as you bite, grind, and chew the meat and bun with your teeth. Unlike the simple, pointed teeth of fish, amphibians, and reptiles, the teeth of mammals vary in size and shape and are specialized to perform specific functions. The chisel-shaped incisors are used for biting and cutting food, whereas the long, pointed canines are adapted for piercing prey and tearing food (FIG. 47-6). The flattened surfaces of the premolars and molars are specialized for crushing and grinding. Each tooth is covered by enamel, the hardest substance in the body (FIG. 47-7). Most of the tooth consists of dentin, which resembles bone in composition and hardness. The pulp cavity, which lies beneath the dentin, is a soft connective tissue containing blood vessels, lymph vessels, and nerves. While food is being mechanically disassembled by the teeth, it is moistened by saliva. As some of the food molecules dissolve,

Incisors Premolars

Molars

(b) Herbivore. Horses and other herbivores have incisors (and sometimes canines) adapted for cutting off bits of vegetation. Canines are absent in some herbivores. The broad, ridged surfaces of the molars are adapted for grinding plant material.

Canines Incisors Premolars

Molars

(c) Omnivore. The teeth of humans and other omnivores are adapted for chewing a variety of foods.

FIGURE 47-6 Teeth and diet

you can taste the food. (Recall from Chapter 43 that taste buds are located on the tongue and other surfaces of the mouth.) Three pairs of salivary glands secrete about a liter of saliva into the mouth cavity each day. Saliva contains an enzyme, salivary amylase, which begins the chemical digestion of starch into sugar.

The pharynx and esophagus conduct food to the stomach After being chewed and fashioned into a lump called a bolus, the bite of food is swallowed—moved through the pharynx into the

Enamel Gum

Crown

Pulp cavity Pulp

Neck

Dentin Cementum Root canal

Root

Spongy bone Nerve Vein

(a) Human lower molar. Sagittal section showing the crown, neck, and root.

Artery

(b) X-ray of a healthy tooth.

FIGURE 47-7 Tooth structure

esophagus. The pharynx, or throat, is a muscular tube that serves as the hallway of both the respiratory system and the digestive system. During swallowing, a small flap of tissue, the epiglottis, closes the opening to the airway. Waves of peristalsis sweep the bolus through the pharynx and esophagus toward the stomach (FIG. 47-8). Circular muscle fibers in the wall of the esophagus contract around the top of the bolus and push it downward. Almost at the same time, longitudinal muscles around the bottom of the bolus and below it contract, which shortens the tube. When the body is in an upright position, gravity helps move the food through the esophagus, but gravity is not essential. Astronauts eat in its absence, and even if you are standing on your head, food will reach your stomach.

Circular muscles contract, constricting passageway and pushing bolus ahead

Food bolus

Longitudinal muscles contract, shortening passageway ahead of the bolus

Relaxed muscle layer

Sphincter closed

Sphincter open o

Stomach

1

Food is mechanically and enzymatically digested in the stomach

Relaxed muscle layer

Esophagus

A bolus is moved through the esophagus by peristaltic contractions.

Stomach

2

When the sphincter (ring of muscle) at the entrance of the stomach opens, food enters the stomach.

The entrance to the large, muscular FIGURE 47-8 Animated Peristalsis stomach is normally closed by a ring of muscle at the lower end of the esophaThe stomach is lined with a simple, columnar epithelium that gus. When a peristaltic wave passes down the esophagus, the muscle secretes large amounts of mucus. Tiny pits mark the entrances to relaxes and the bolus enters the stomach (FIG. 47-9). When empty, the millions of gastric glands, which extend deep into the stomach the stomach is collapsed and shaped almost like a hot dog. Folds wall. Parietal cells in the gastric glands secrete hydrochloric acid of the stomach wall, called rugae, give the inner lining a wrinkled and intrinsic factor, a substance needed for adequate absorption of appearance. As food enters, the rugae gradually smooth out, which vitamin B12. Chief cells in the gastric glands secrete pepsinogen, expands the capacity of the stomach to more than a liter.

KEY POINT

The structure of the stomach is adapted for its function of digesting food; gastric glands in its mucosa secrete digestive juices, and three layers of muscle in the stomach wall are important in mechanically digesting food.

Visceral peritoneum Esophagus Sphincter Chief cell

Circular muscle layer

Parietal cell

Longitudinal muscle layer Oblique muscle layer Pyloric sphincter

(a)

Nuclei

Duodenum

Surface epithelium

Rugae

Chief cells Openings into gastric glands

Parietal cells Gastric glands

Gastric glands

Epithelium

Lymph nodule

(b)

Gastric mucosa

Gastric glands

FIGURE 47-9 Structure of the stomach The stomach is adapted for mechanically and enzymatically digesting food. (a) The wall of the stomach has been progressively removed to show muscle layers and rugae. (b) Stomach lining and gastric glands.

an inactive enzyme precursor. When it comes in contact with the acidic gastric juice in the stomach, pepsinogen is converted to pepsin, the main digestive enzyme of the stomach. Pepsin hydrolyzes proteins, converting them to short polypeptides. Several protective mechanisms prevent the gastric juice from digesting the wall of the stomach. Cells of the gastric mucosa secrete alkaline mucus that coats the stomach wall and neutralizes the acidity of the gastric juice along the lining. In addition, the epithelial cells of the lining fit tightly together, preventing gastric juice from leaking between them and into the tissue beneath. If some of the epithelial cells become damaged, they are quickly replaced. In fact, about a half million of these cells are shed and replaced every minute! Sometimes, these protective mechanisms malfunction and part of the stomach lining is digested, leaving an open sore, or peptic ulcer. Such ulcers also occur in the duodenum and sometimes in

the lower part of the esophagus. The bacterium Helicobacter pylori has been implicated as a causative factor in ulcers (see Fig. 25-13). Helicobacter pylori infects the mucus-secreting cells of the stomach lining, decreasing the protective mucus, which can lead to peptic ulcers or cancer. This type of infection responds to antibiotic therapy. What changes occur in a bite of hamburger during its 3- to 4-hour stay in the stomach? The stomach churns and chemically degrades the food so that it assumes the consistency of a thick soup; this partially digested food is called chyme. Protein digestion then degrades much of the hamburger protein to polypeptides. Digestion of the starch in the bun to small polysaccharides and maltose continues until salivary amylase is inactivated by the acidic pH of the stomach. Over a period of several hours, peristaltic waves release the chyme in spurts through the stomach exit, the pylorus, and into the small intestine.

Most enzymatic digestion takes place in the small intestine Digestion of food is completed in the small intestine, and nutrients are absorbed through its wall. The small intestine, which is 5 to 6 m (about 17 ft) in length, has three regions: the duodenum, jejunum, and ileum. Most chemical digestion takes place in the duodenum, the first portion of the small intestine, not in the stomach. Bile from the liver and enzymes from the

The digestive and absorptive surface area of the small intestine is vastly increased by millions of fingerlike villi; the surface area is further increased by microvilli on the epithelial cells of the villi. G. Shih–R. Kessel/Visuals Unlimited

KEY POINT

pancreas are released into the duodenum and act on the chyme. Then enzymes produced by the epithelial cells lining the duodenum catalyze the final steps in the digestion of the major types of nutrients. The lining of the small intestine appears velvety because of its millions of tiny fingerlike projections, the intestinal villi (sing., villus) (FIG. 47-10). The villi increase the surface area of the small intestine for digestion and absorption of nutrients. The intestinal surface is further expanded by microvilli, projections of the

(a) SEM of a cross section of the small intestine.

50 μm Courtesy of J. D. Hoskings, W. G. Henk, and Y. Z. Abdelbaki, American Journal of Veterinary Research, 1982, Vol. 43, No. 10, pp. 1715–1720

Villi

Epithelial cells lining villus

Capillary network Nerve fiber Mucosa

Openings of intestinal glands Goblet cells Intestinal glands

Submucosa Muscle layer

Lacteal Lymph vessel Visceral peritoneum

(b) Villi. Enlarged view of a small portion of the intestinal wall. Some of the villi have been opened to show the blood and lymph vessels within.

FIGURE 47-10 Villi and microvilli

1 μm

(c) Microvilli. SEM of the surface of an epithelial cell from the lining of the small intestine, showing microvilli. The epithelium has been cut vertically, allowing the microvilli to be viewed from the side as well as from above.

plasma membrane of the simple columnar epithelial cells of the villi. About 600 microvilli protrude from the exposed surface of each cell, giving the epithelial lining a fuzzy appearance when viewed with the light microscope. If the intestinal lining were smooth, like the inside of a water pipe, food would zip right through the intestine, and many valuable nutrients would not be digested or absorbed. Folds in the wall of the intestine, the villi, and microvilli together increase the surface area of the small intestine by about 600 times. If we could unfold and spread out the lining of the small intestine of an adult human, its surface would approximate the size of a tennis court.

Stomach Inferior vena cava Right lobe of liver Right hepatic duct

Pancreas

The liver secretes bile The liver, the largest internal organ and also one of the most complex organs in the body, lies in the upper-right part of the abdomen just under the diaphragm (FIG. 47-11). The liver secretes bile, which mechanically digests fats by a detergent-like action (discussed in a later section). Bile consists of water, bile salts, bile pigments, cholesterol, salts, and lecithin (a phospholipid). Because it contains no digestive enzymes, bile does not enzymatically digest food. The pear-shaped gallbladder stores and concentrates bile and releases it into the duodenum as needed. A single liver cell can carry on more than 500 separate, specialized metabolic activities! The liver performs these vital functions: 1. Secretes bile that mechanically digests fats 2. Helps maintain homeostasis by removing or adding nutrients to the blood 3. Converts excess glucose to glycogen and stores it 4. Converts excess amino acids to fatty acids and urea 5. Stores iron and certain vitamins 6. Detoxifies alcohol and other drugs and poisons

The pancreas secretes digestive enzymes The pancreas is an elongated gland that secretes both digestive enzymes and hormones that help regulate the level of glucose in the blood. The cells that line the pancreatic ducts secrete an alkaline solution rich in bicarbonate ions. This pancreatic juice neutralizes the stomach acid in the duodenum and provides the optimal pH for action of the pancreatic enzymes. Pancreatic enzymes include trypsin and chymotrypsin, which digest polypeptides to dipeptides; pancreatic lipase, which degrades fats; pancreatic amylase, which breaks down almost all types of complex carbohydrates, except cellulose, to disaccharides; and ribonuclease and deoxyribonuclease, which split the nucleic acids ribonucleic acid (RNA) and deoxyribonucleic acid (DNA) to free nucleotides.

Nutrients are digested as they move through the digestive tract Chyme moves through the digestive tract by peristalsis, mixing contractions, and motions of the villi. As nutrients in the chyme move through the small intestine, they come into contact with enzymes that digest them (TABLE 47-1).

Common bile duct

Hepatic portal vein Pancreatic duct

Gallbladder

Duodenum

FIGURE 47-11 The liver and pancreas The gallbladder stores bile from the liver. Note the ducts that conduct bile to the gallbladder and the duodenum. The stomach has been displaced to expose the pancreas, which secretes several digestive enzymes into the duodenum.

Carbohydrates are digested to monosaccharides Polysaccharides, such as starch and glycogen, are important components of the food ingested by most animals. The glucose units of these large molecules are connected by glycosidic bonds linking carbon 4 (or 6) of one glucose molecule with carbon 1 of the adjacent glucose molecule. These bonds are hydrolyzed by amylases that digest polysaccharides to the disaccharide maltose. Although amylase can split the a-glycosidic linkages present in starch and glycogen, it cannot split the b-glycosidic linkages present in cellulose (see Figs. 3-9 and 3-10). Amylase cannot split the bond between the two glucose units of maltose. Enzymes produced by the cells lining the small intestine break down disaccharides such as maltose to monosaccharides. Maltase, for example, splits maltose into two glucose molecules (see Fig. 3-8a). Hydrolysis occurs while the disaccharides are being absorbed through the epithelium of the small intestine. Proteins are digested to amino acids Several proteolytic enzymes are secreted into the digestive tract. Each breaks peptide bonds at one or more specific locations in a polypeptide chain. Trypsin, secreted in an inactive form by the pancreas, is activated by an enzyme called enterokinase. The trypsin then activates chymotrypsin and carboxypeptidase, as well as additional trypsin. Pepsin, trypsin, and chymotrypsin break certain internal peptide bonds of proteins and polypeptides. Carboxypeptidase removes amino acids with free terminal carboxyl groups from the end of polypeptide chains. Dipeptidases released by the duodenum then split the small peptides to amino acids.

TABLE 47-1

Summary of Digestion Carbohydrates

Mouth

Proteins

Lipids

Polysaccharides ¡

Salivary amylase

Maltose and small polysaccharides Stomach

Protein ¡

Action continues until acidic pH inactivates salivary amylase

Pepsin

Short polypeptides Small intestine

Undigested polysaccharides

Polypeptides

Bile salts

Maltose and other disaccharides

Short polypeptides and peptides

Emulsified fat droplets

¡

¡

¡

Maltase, sucrase, lactase

Monosaccharides

Fats are digested to fatty acids and monoacylglycerols Lipids are usually ingested as large masses of triacylglycerols (also called triglycerides). They are digested mainly within the duodenum by pancreatic lipase. Like many other proteins, lipase is water soluble, but its substrates are not. Thus, the enzyme can attack only the fat molecules at the surface of a mass of fat. Bile salts act like detergents, reducing the surface tension of fats. Their action, called emulsification, breaks large masses of fat into smaller droplets. Emulsification greatly increases the surface area of fat exposed to the action of pancreatic lipase and so increases the rate of lipid digestion. Conditions in the intestine are usually not optimal for the complete hydrolysis of lipids to glycerol and fatty acids. Consequently, the products of lipid digestion include monoacylglycerols (monoglycerides) and diacylglycerols (diglycerides) as well as glycerol and fatty acids. Undigested triacylglycerols also remain, some of which are absorbed without digestion.

TABLE 47-2

Trypsin, chymotrypsin

¡

¡

¡

Pancreatic amylase

Glob of fat

Carboxypeptidase, peptidases, dipeptidases

Amino acids

Pancreatic lipase

Fatty acids and glycerol

Nerves and hormones regulate digestion Most digestive enzymes are produced only when food is present in the digestive tract. Salivary gland secretion is controlled entirely by the nervous system, but secretion of other digestive juices is regulated by both nerves and hormones. The wall of the digestive tract contains dense networks of neurons. This so-called enteric nervous system continues to regulate many motor and secretory activities of the digestive system even if sympathetic and parasympathetic nerves to these organs are cut. Many neuropeptides present in the brain are also released by neurons in the digestive tract and help regulate digestion. For example, substance P stimulates smooth muscle contraction of the digestive tract, and enkephalin inhibits it (see Table 41-2). Several hormones, including gastrin, secretin, cholecystokinin (CCK), and glucose-dependent insulinotropic peptide (GIP), help regulate the digestive system (TABLE 47-2). These hormones are polypeptides secreted by endocrine cells in the mucosa

Some Hormones That Regulate Digestion

Hormone

Source

Target Tissue

Actions

Factors That Stimulate Release

Gastrin

Stomach (mucosa)

Stomach (gastric glands)

Stimulates gastric glands to secrete pepsinogen and HCl; stimulates gastric motility

Distention of the stomach by food; certain substances such as partially digested proteins and caffeine

Secretin

Duodenum (mucosa)

Pancreas

Signals secretion of sodium bicarbonate Stimulates bile secretion Inhibits gastric secretion

Acidic chyme acting on mucosa of duodenum

Presence of fatty acids and partially digested proteins in duodenum

Gallbladder Brain

Stimulates release of digestive enzymes Stimulates emptying of bile Signals satiety

Pancreas

Stimulates insulin secretion

Presence of glucose in duodenum

Liver Stomach Cholecystokinin (CCK)

Glucose-dependent insulinotropic peptide (GIP)

Duodenum (mucosa)

Duodenum (mucosa)

Pancreas

of certain regions of the digestive tract. Investigators are studying several other messenger peptides that are important in regulating digestive activity. As an example of the regulation of the digestive system, consider the secretion of gastric juice. Seeing, smelling, tasting, or even thinking about food causes the brain to send neural signals to the gastric glands in the stomach. These signals stimulate these glands to secrete gastric juice. In addition, when food distends the stomach, stretch receptors send neural messages to the medulla. The medulla then sends messages to endocrine cells in the stomach wall that secrete gastrin. This hormone is absorbed into the blood; it stimulates the stomach to release gastric juice and also stimulates gastric emptying and intestinal motility.

Absorption takes place mainly through the villi of the small intestine Only a few substances—water, simple sugars, salts, alcohol, and certain drugs—are small enough to be absorbed through the stomach wall. Absorption of nutrients is primarily the job of the intestinal villi. As illustrated in Figure 47-10, the wall of a villus is a single layer of epithelial cells. Inside each villus is a network of capillaries and a central lymph vessel, called a lacteal. To reach the blood (or lymph), a nutrient molecule must pass through an epithelial cell lining the intestine and through a cell lining a blood or lymph vessel. Absorption occurs by a combination of simple diffusion, facilitated diffusion, and active transport. Because glucose and amino acids cannot diffuse through the intestinal lining, they must be absorbed by active transport. Absorption of these nutrients is coupled with the active transport of sodium (see Chapter 5). Fructose is absorbed by facilitated diffusion. Amino acids and glucose are transported directly to the liver by the hepatic portal vein. In the liver this vein divides into a vast network of tiny blood sinuses, vessels similar to capillaries. As the nutrient-rich blood moves slowly through the liver, nutrients and certain toxic substances are removed from the circulation. The products of lipid digestion are absorbed by a different process and different route. After free fatty acids and monoacylglycerols enter an epithelial cell in the intestinal lining, they are reassembled as triacylglycerols in the smooth endoplasmic reticulum. The triacylglycerols, along with absorbed cholesterol and phospholipids, are packaged into protein-covered fat droplets, called chylomicrons. After they are released into the interstitial fluid, chylomicrons enter the lacteal (lymph vessel) of the villus. Chylomicrons are transported in the lymph to the subclavian veins, where the lymph and its contents enter the blood. About 90% of absorbed fat enters the blood circulation in this indirect way. The rest, mainly shortchain fatty acids such as those in butter, are absorbed directly into the blood. After a meal rich in fats, the great number of chylomicrons in the blood may give the plasma a turbid, milky appearance for a few hours. Chylomicrons transport fats to the liver and other tissues. Most of the nutrients in the chyme are absorbed by the time it reaches the end of the small intestine. What is left (mainly waste) passes through a sphincter, the ileocecal valve, into the large intestine.

The large intestine eliminates waste Undigested material, such as the cellulose of plant foods, along with unabsorbed chyme, passes into the large intestine (see Fig. 47-4). Although only about 1.3 m (about 4 ft) long, this part of the digestive tract is referred to as “large” because its diameter is greater than that of the small intestine. The small intestine joins the large intestine about 7 cm (2.8 in) from the end of the large intestine, forming a blind pouch, the cecum. The vermiform appendix projects from the end of the cecum. (Appendicitis is an inflammation of the appendix.) Herbivores such as rabbits have a large, functional cecum that holds food while bacteria digest its cellulose. In humans, the functions of the cecum and appendix are not known, and these structures are generally considered vestigial organs. From the cecum to the rectum (the last portion of the large intestine), the large intestine is known as the colon. The regions of the large intestine are the cecum; ascending colon; transverse colon; descending colon; sigmoid colon; rectum; and anus, the opening for the elimination of wastes. As chyme passes slowly through the large intestine, water and sodium are absorbed from it, and it gradually assumes the consistency of normal feces. Bacteria inhabiting the large intestine are nourished by the last remnants of the meal; in exchange, they benefit their host by producing vitamin K and certain B vitamins that can be absorbed and used. A distinction should be made between elimination and excretion. Elimination is the process of getting rid of digestive wastes— materials that have not been absorbed from the digestive tract and did not participate in metabolic activities. In contrast, excretion is the process of getting rid of metabolic wastes, which in mammals is mainly the function of the kidneys and lungs. The large intestine, however, does excrete bile pigments. When chyme passes through the intestine too rapidly, defecation (expulsion of feces) becomes more frequent, and the feces are watery. This condition, called diarrhea, may be caused by anxiety, certain foods, or pathogens that irritate the intestinal lining. Prolonged diarrhea results in loss of water and salts and leads to dehydration—a serious condition, especially in infants. Constipation results when chyme passes through the intestine too slowly. Because more water than usual is removed from the chyme, the feces may be hard and dry. Constipation is often caused by a diet deficient in fiber. Colorectal cancer occurs in the colon or rectum (FIG. 47-12). In Western countries, colorectal cancer is the third most common cancer, and in the United States, it is the second leading cancer killer. Several factors contribute to the risk of developing colon cancer, including family history, certain genetic factors, smoking, physical inactivity, heavy use of alcohol, and obesity. A diet high in red meat and processed meat and low in fresh fruit, vegetables, poultry, and fish appears to increase the risk.

Review ■

■ ■ ■

Imagine that you have just taken a bite of a chicken sandwich. List in sequence the structures through which it passes in its journey through the digestive system. What happens in each structure? What is the function of villi? How do they function? What is the function of the pancreas? What are chylomicrons?

Transverse colon Ascending colon

Recall from Chapter 1 that metabolism includes all the chemical processes that take place in the body. Metabolism includes anabolic and catabolic processes. Anabolism includes the synthetic aspects of metabolism, such as the production of proteins and nucleic acids. Catabolism includes breakdown processes, such as hydrolysis. Nutritionists measure the energy value of food in kilocalories, or simply Calories. A Calorie, spelled with a capital C, denotes a kilocalorie (kcal), defined as the amount of heat required to raise the temperature of a kilogram of water 1 degree Celsius.

Carbohydrates provide energy Cancer

SPL / Photo Researchers, Inc.

Descending colon

FIGURE 47-12 Colon cancer In this radiographic view of the large intestine, cancer is evident as a mass that projects into the lumen of the colon. The large intestine has been filled with a suspension of barium sulfate, which makes irregularities in the wall visible.

47.3 REQUIRED NUTRIENTS ■ ■ LEARNING OBJECTIVES 6 Summarize the nutritional requirements for dietary carbohydrates,

7

lipids, and proteins, and trace the fate of glucose, lipids, and amino acids after their absorption. Describe the nutritional functions of vitamins, minerals, and phytochemicals.

Adequate amounts of essential nutrients are necessary for metabolic processes. Animals require carbohydrates, lipids, proteins, vitamins, and minerals. Nutritionists are also investigating plant compounds called phytochemicals that are important in nutrition. Although not considered a nutrient in a strict sense, water is a necessary dietary component. Enough fluid must be ingested to replace fluid lost in urine, sweat, feces, and breath. Nutritionists consider new data and continually re-examine the relative amounts of various types of nutrients recommended for a healthy diet. According to the U.S. Department of Agriculture’s Dietary Guidelines for Americans, 45% to 65% of our calories should come from carbohydrates, 20% to 35% from fat, and 10% to 35% from protein.

Sugars and starches are important sources of energy in the human diet. Most carbohydrates are ingested in the form of starch and cellulose, both polysaccharides composed of long chains of glucose subunits. (You may want to review the discussion of carbohydrates in Chapter 3.) Nutritionists refer to polysaccharides as “complex carbohydrates.” Foods rich in complex carbohydrates include rice, potatoes, corn, and other cereal grains. When we eat an excess of carbohydrate-rich food, the liver cells become fully packed with glycogen and convert excess glucose to fatty acids and glycerol. Liver cells convert these compounds to triacylglycerols and send them to the fat depots of the body for storage. Refined carbohydrates, such as white bread and white rice, are unhealthy because the refining process removes fiber and many vitamins and minerals. The refining process also produces a form of starch that the digestive system rapidly breaks down to glucose. The resulting rapid increase in glucose concentration in the blood stimulates the pancreas to release a large amount of insulin. This hormone lowers blood glucose level by signaling the liver and muscles to remove glucose from the blood. When glucose and insulin levels are high, triacylglycerol levels increase and high-density lipoprotein (HDL; the good cholesterol) concentration decreases. These metabolic events can lead to cardiovascular disease and increase the risk of type 2 diabetes (discussed in Chapter 49). Dietary fiber decreases cholesterol concentration in the blood and is associated with a lower risk for cardiovascular disease and diabetes. Fiber is mainly a mixture of cellulose and other indigestible carbohydrates. We obtain dietary fiber by eating fruits, vegetables, and whole grains. The U.S. diet is low in fiber because of low intake of fruits and vegetables and use of refined flour. Increasing fiber in the diet has a variety of health benefits. Fiber stimulates the feeling of being satisfied (satiety) after food intake and thus is useful in treating obesity.

Lipids provide energy and are used to make biological molecules Cells use ingested lipids to provide energy and to make a variety of lipid compounds, such as components of cell membranes, steroid hormones, and bile salts. We ingest about 98% of our dietary lipids in the form of triacylglycerols (triglycerides). (Recall from Chapter 3 that a triacylglycerol is a glycerol molecule chemically combined with three fatty acids; see Fig. 3-12b.) Triacylglycerols may be saturated, that is, fully loaded with hydrogen atoms, or their fatty

In general, animal foods are rich in both saturated fats and acids may be monounsaturated (containing one double bond) or cholesterol, whereas most plant foods contain unsaturated fats and polyunsaturated (containing two or more double bonds). no cholesterol. Butter contains mainly saturated fats. Commonly Three polyunsaturated fatty acids (linoleic, linolenic, and used polyunsaturated vegetable oils are corn, soybean, and safarachidonic acids) are essential fatty acids that humans must obflower oils. Monounsaturated fats raise HDL level, as does stearic tain from food. Given these and sufficient nonlipid nutrients, the acid, the saturated fat in chocolate. Olive, canola, and peanut body can make all the lipid compounds (including fats, cholesoils contain large amounts of monounsaturated fats. A few plant terol, phospholipids, and prostaglandins) that it needs. The averoils, including palm oil and coconut oil, are high in saturated fats. age U.S. diet provides far more cholesterol than the recommended Omega-3 fatty acids (found in fish and some plant oils) decrease daily maximum of 300 mg. High-cholesterol sources include egg LDL levels and play other protective roles in decreasing the risk for yolks, butter, and meat. coronary heart disease. How are lipids transported? Recall that chylomicrons transport A diet high in saturated fats and cholesterol raises the blood lipids from the intestine to the liver and to other tissues. As chylocholesterol level by as much as 25%. Because polyunsaturated fats microns circulate in the blood, the enzyme lipoprotein lipase decrease the blood cholesterol level, many people now cook with breaks down triacylglycerols. The fatty acids and glycerol can then vegetable oils rather than with butter and lard, drink skim milk be taken up by the cells. What is left of the chylomicron, a remnant rather than whole milk, and eat low-fat ice cream. Some use margamade up mainly of cholesterol and protein, is taken up by the liver. rine instead of butter. However, during production of many margaLiver cells repackage cholesterol and triacylglycerols. These rines, vegetable oils are partially hydrogenated (some of the carbons lipids are bound to proteins and transported as large molecular accept hydrogen to become fully saturated). During hydrogenation, complexes, called lipoproteins. Some plasma cholesterol is transsome double bonds are transformed from a cis arrangement to a ported by high-density lipoproteins, or HDLs (“good” cholestrans arrangement (see Fig. 3-3b), forming trans fatty acids; the terol), but most is transported by low-density lipoproteins, or harder the margarine, the higher the trans fatty acid content. Health LDLs (“bad” cholesterol). LDLs deliver cholesterol to the cells. For cells to take in LDLs, a protein (apolipoprotein B) on the LDL surface must bind with a protein LDL receptor on the plasma membrane (see Fig. 5-23). After binding, Fat cell Fat the LDL enters the cell, and its cholesterol and other components are used. When cholesterol levels are high, HDLs Fatty acids collect the excess cholesterol and transport it to the liver. + HDLs decrease risk for cardiovascular disease. Glycerol When needed, stored fats are hydrolyzed to fatty acids and released into the blood. Before cells can use these Transported in blood fatty acids as fuel, they are broken down into smaller comLiver pounds and combined with coenzyme A to form molecules Fatty acid + Glycerol of acetyl coenzyme A (acetyl CoA; FIG. 47-13). Acetyl CoA Enters enters the citric acid cycle (see Chapter 8). The conversion G3P cellular Used Acetyl CoA of fatty acids to acetyl CoA is accomplished in the liver by respiration to make a process known as b-oxidation. triacylglycerols Glucose For transport to the cells, acetyl coenzyme A is conConverted verted into one of three types of ketone bodies (four-carbon to other lipids Ketone bodies ketones). Normally, the level of ketone bodies in the blood Enters is low, but in certain abnormal conditions, such as starvacellular respiration tion and diabetes mellitus, fat metabolism is tremendously increased. Ketone bodies are then produced so rapidly Other that their level in the blood becomes excessive, which may Pyruvate cells make the blood too acidic. Such disrupted pH balance can or acetyl CoA lead to death. What is the relationship between fat and cholesterol intake and cardiovascular disease? Lipids play a key role in Converted Enters to other the development of atherosclerosis, a progressive disease cellular lipids respiration in which the arteries become clogged with fatty material (see Chapter 44). LDLs are the main source of the cholesterol that builds up in the walls of arteries. The type of fat consumed, as well as other dietary and lifestyle factors, is important. You can promote a healthy proportion of HDL to LDL by exercising regularly, following a healthy diet, FIGURE 47-13 How the body uses fat maintaining appropriate body weight (obesity raises LDL The liver converts glycerol and fatty acids to compounds that are used as fuel in cellular respiration. Recall from Chapter 8 that G3P is glyceraldehyde-3-phosphate. and triacylglycerol levels), and not smoking cigarettes.

concerns have been raised because trans fatty acids increase LDL cholesterol in the blood and lower HDL. Many processed foods, including cakes, doughnuts, cookies, crackers, and potato chips, contain trans fatty acids. Although some fats, such as trans fatty acids and saturated fats, are unhealthy, others, such as omega-3 fatty acids and monounsaturated fat, promote health. The Inuits in Greenland eat diets extremely high in fat. However, because their dietary fat is rich in omega-3 fatty acids, they have a low incidence of heart disease.

Proteins serve as enzymes and as structural components of cells

AMINO ACIDS Catabolism Excess amino acids Deamination Liver

Anabolism Structural proteins, hemoglobin, myosin, actin, enzymes, plasma proteins

NH3 (ammonia) + α-keto acids Fat Urea

Pyruvate, α-ketoglutarate

Enter cellular respiration

Proteins are essential building blocks of cells, serve as enAcetyl CoA zymes, and are used to make essential compounds such as receptors, hemoglobin and myosin. Protein consumption Ketone bodies is an index of a country’s (or an individual’s) economic status because high-quality protein tends to be the most expensive and least available of the nutrients. In many parts To kidneys of the world, protein poverty is one of the most pressing Acetyl CoA health problems. Ingested proteins are degraded in the digestive tract to Fatty acids + glycerol small peptides and amino acids. Of the 20 or so amino acids important in nutrition, approximately 9 (10 in children) cannot be synthesized by humans at all, or at least not in Triacylglycerol sufficient quantities to meet the body’s needs. The diet must provide these essential amino acids (see Chapter 3). Storage in fat cells Complete proteins, those that contain the most appropriate distribution of amino acids for human nutrition, are found in fish, meat, nuts, eggs, and milk. Some foods, such FIGURE 47-14 How the body uses protein as gelatin and legumes (soybeans, beans, peas, peanuts), The liver plays a central role in protein metabolism. Deamination of amino acids and contain a high proportion of protein. However, they either conversion of the amino groups to urea take place there. In addition, many proteins do not contain all the essential amino acids, or they do are synthesized in the liver. Note that you can gain weight on a high-protein diet not contain them in proper nutritional proportions. Most because excess amino acids can be converted to fat. plant proteins are deficient in one or more essential amino acids. The healthiest sources of protein are fish, chicken, mins. Fruit and vegetables are rich sources of vitamins. TABLE 47-3 nuts, and legumes. provides the sources, functions, and consequences of deficiency Amino acids circulating in the blood are taken up by cells and for most of the vitamins. used for the synthesis of proteins. The liver removes excess amino Health professionals debate the advisability of taking large acids from the circulation. Liver cells deaminate amino acids, that amounts of certain specific vitamins, such as vitamin C to prevent is, remove the amino group (FIG. 47-14). During deamination, ammonia is produced from the amino group. Ammonia, which is toxic colds or vitamin E to protect against vascular disease. Some studat high concentrations, is converted to urea and excreted from the ies suggest that vitamin A (found in yellow and green vegetables) body. The remaining carbon chain of the amino acid (called a keto and vitamin C (found in citrus fruit and tomatoes) help protect acid) may be converted into carbohydrate or lipid and used as fuel against certain forms of cancer. We do not yet understand all the or stored. This is why people who eat high-protein diets can gain biochemical roles played by vitamins or the interactions among weight if they eat too much. various vitamins and other nutrients. We do know that, like vitamin deficiency, large overdoses of vitamins can be harmful. Moderate overdoses of the B and C vitamins are excreted in the urine, Vitamins are organic compounds but surpluses of the fat-soluble vitamins are not easily excreted and essential for normal metabolism can accumulate to harmful levels. Vitamins are organic compounds required in the diet in relatively small amounts for normal biochemical functioning. Many are Minerals are inorganic nutrients components of coenzymes (see Chapter 7). Nutritionists divide Minerals are inorganic nutrients ingested in the form of salts disvitamins into two main groups. Fat-soluble vitamins include vitasolved in food and water (TABLE 47-4). We need certain minermins A, D, E, and K. Water-soluble vitamins are the B and C vita-

TABLE 47-3

The Vitamins

Vitamins and U.S. RDA*

Actions

Effect of Deficiency

Sources

Vitamin A, retinol 5000 IU†

Converted to retinal; essential for normal vision; essential for normal growth and differentiation of cells; reproduction; immunity

Growth retardation; night blindness; worldwide millions of children are at risk of blindness from vitamin A deficiency

Liver, fortified milk, orange and green vegetables such as carrots and broccoli; orange fruits such as cantaloupe

Vitamin D, calciferol 400 IU

Promotes calcium and phosphorus absorption from digestive tract; essential to normal growth and maintenance of bone

Weak bones; bone deformities; rickets in children, osteomalacia in adults

Fish oils, egg yolk, fortified milk, butter, margarine, salmon

Vitamin E, tocopherols 30 IU

Antioxidant; protects unsaturated fatty acids and cell membranes

Increased catabolism of unsaturated fatty acids so that not enough are available for maintenance of cell membranes; prevention of normal growth; nerve damage

Polyunsaturated plant oils, nuts, leafy greens, whole-grain products

Vitamin K, about 80 mcg‡

Synthesis of blood-clotting proteins and proteins important in bone production

Prolonged blood-clotting time

Normally supplied by intestinal bacteria; leafy greens, legumes

Collagen synthesis; antioxidant; needed for synthesis of some hormones and neurotransmitters; important in immune function

Scurvy (wounds heal very slowly, and scars become weak and split open; capillaries become fragile; bone does not grow or heal properly); immune system suppressed

Citrus fruit, strawberries, tomatoes, leafy vegetables, cabbage

Active form is a coenzyme in many enzyme systems; important in carbohydrate and amino acid metabolism

Beriberi (weakened heart muscle, enlarged right side of heart, nervous sytem and digestive tract disorders); common in alcoholics

Liver, yeast, whole and enriched grains, meat, green leafy vegetables

Vitamin B2, riboflavin 1.7 mg

Used to make coenzymes (e.g., FAD) essential in cellular respiration

Dermatitis, inflammation and cracking at corners of mouth; confusion

Liver, milk, eggs, green leafy vegetables, enriched grains

Niacin, 20 mg

Component of important coenzymes (NAD+ and NADP+); essential to cellular respiration

Pellagra (dermatitis, diarrhea, mental symptoms, muscular weakness, fatigue)

Liver, chicken, tuna, milk, green leafy vegetables, enriched grains

Vitamin B6, pyridoxine 2 mg

Derivative is coenzyme in many reactions in amino acid metabolism

Dermatitis, digestive tract disturbances; convulsions

Meat, whole grains, legumes, green leafy vegetables

Pantothenic acid, 10 mg

Constituent of coenzyme A (important in cellular metabolism)

Deficiency extremely rare

Meat, whole grains, legumes

Folate, 400 mcg

Coenzyme needed for nucleic acid synthesis and for maturation of red blood cells

A type of anemia; certain birth defects; increased risk of cardiovascular disease; deficiency in alcoholics, smokers, and pregnant women

Liver, legumes, dark green leafy vegetables, orange juice, grains

Biotin, 30 mcg

Coenzyme important in metabolism

Deficiency uncommon; depression, skin irritation, seizures

Produced by intestinal bacteria; liver, chocolate, egg yolk

Vitamin B12, 2.4 mcg

Coenzyme important in metabolism; contains cobalt

A type of anemia

Liver, meat, fish, eggs, cheese

Fat soluble

Water soluble Vitamin C, ascorbic acid 60 mg

B-complex vitamins Vitamin B1, thiamine 1.5 mg

*RDA is the recommended dietary allowance, established by the Food and Nutrition Board of the National Research Council, to maintain good nutrition for healthy adults. †

International Unit: the amount that produces a specific biological effect and is internationally accepted as a measure of the activity of the substance.

‡

mcg = micrograms.

als, including sodium, chloride, potassium, calcium, phosphorus, magnesium, manganese, and sulfur, in amounts of 100 mg or more daily. These are referred to as major minerals. Several others, such as iron, copper, iodide, fluoride, and selenium, are trace elements, minerals that we require in amounts less than 100 mg per day. Minerals are necessary components of body tissues and fluids. Salt content (about 0.9% in plasma) is vital in maintaining the fluid balance of the body, and because salts are lost from the body daily in sweat, urine, and feces, they must be replaced by dietary intake. Sodium chloride (common table salt) is the salt needed in largest quantity in blood and other body fluids. Deficiency results

in dehydration. Iron is the mineral most often deficient in the diet. In fact, iron deficiency is one of the most widespread nutritional problems in the world.

Antioxidants protect against oxidants Normal cell processes that require oxygen produce oxidants, highly reactive molecules such as free radicals, peroxides, and superoxides. Free radicals, molecules or ions with one or more unpaired electrons, are also generated by ionizing radiation, tobacco smoke, and other forms of air pollution. Oxidants damage DNA,

TABLE 47-4

Some Important Minerals and Their Functions

Mineral

Functions

Sources; Comments

Calcium

Main mineral of bones and teeth; essential for normal blood clotting, muscle function, nerve function, and regulation of cell activities

Milk and other dairy products, fish, green leafy vegetables; bones serve as calcium reservoir

Phosphorus

Performs more functions than any other mineral; structural component of bone; component of ATP, DNA, RNA, and phospholipids

Meat, dairy products, cereals

Sulfur

Component of many proteins and vitamins

High-protein foods such as meat, fish, legumes, nuts

Potassium

Principal positive ion within cells; important in muscle contraction and nerve function

Meat, milk, fruit, vegetables, grains

Sodium

Principal positive ion in interstitial fluid; important in fluid balance; neural transmission

Many foods, table salt; too much ingested in average American diet; excessive amounts may contribute to high blood pressure

Chloride

Principal negative ion of interstitial fluid; important in fluid balance and in acid–base balance

Many foods, table salt

Magnesium

Needed for normal muscle and nerve function

Nuts, whole grains, green leafy vegetables, seafood, chocolate

Copper

Component of several enzymes; essential for hemoglobin synthesis

Liver, eggs, fish, whole wheat flour, beans

Iodide

Component of thyroid hormones (hormones that increase metabolic rate); deficiency results in goiter (abnormal enlargement of thyroid gland)

Seafood, iodized salt, vegetables grown in iodine-rich soils

Manganese

Activates many enzymes

Whole-grain cereals, nuts, leafy green vegetables; poorly absorbed from intestine

Iron

Component of hemoglobin, myoglobin, important respiratory enzymes (cytochromes), and other enzymes essential to oxygen transport and cellular respiration; deficiency results in anemia and may impair cognitive function

Mineral most likely to be deficient in diet. Good sources: meat (especially liver), fish, nuts, egg yolk, legumes, dried fruit

Fluoride

Component of bones and teeth; makes teeth resistant to decay; excess causes tooth mottling

Fish; in areas where it does not occur naturally, fluoride may be added to municipal water supplies (fluoridation)

Zinc

Cofactor for at least 70 enzymes; helps regulate synthesis of certain proteins; needed for growth and repair of tissues; deficiency associated with loss of appetite, depressed immune function, retarded growth in children

Meat, fish, milk, yogurt, grains, vegetables

Selenium

Antioxidant (cofactor for a peroxidase that breaks down peroxides); appears to protect against prostate cancer in men

Seafood, eggs, meat, whole grains

proteins, and unsaturated fatty acids by snatching electrons. Damage to DNA causes mutations that may lead to cancer, and injury to unsaturated fatty acids can damage cell membranes. Free radicals are thought to contribute to atherosclerosis by causing oxidation of LDL cholesterol. Oxidative damage to the body over many years contributes to the aging process. Cells have antioxidants that destroy free radicals and other reactive molecules. Antioxidants in tissues include certain enzymes, for example, catalase and peroxidase. Their action requires minerals such as selenium, zinc, manganese, and copper. Certain vitamins—vitamin C, vitamin E, and vitamin A—have strong antioxidant activity. Vitamins A and E protect cell membranes from free radicals. In addition, a variety of phytochemicals are potent antioxidants (discussed in next section).

Nutritionists recommend that we increase the antioxidants in our diet by eating fruits, vegetables, and other foods high in antioxidants. We do not yet know whether antioxidant supplements are useful.

Phytochemicals play important roles in maintaining health A diet that includes all the essential nutrients does not provide the same health benefits as one also rich in fruit and vegetables. The missing ingredients are phytochemicals, plant compounds that promote health. Many phytochemicals are antioxidants. For example, lycopenes, responsible for the red color of tomatoes, are powerful antioxidants. Flavonoids, found in cocoa, especially in dark choco-

late, are antioxidants that also appear to lower blood pressure, lower cholesterol, and improve cardiovascular health in other ways. Nutritionists have begun to intensively investigate phytochemicals. Some important classes of phytochemicals are carotenoids (which include vitamin A and lycopene) in deeply pigmented fruits and vegetables such as carrots and tomatoes; flavonoids in berries, herbs, vegetables, barley, soy, black and green tea, and cocoa; isothiocyanates and indoles in broccoli and other cruciferous vegetables; and organosulfur compounds in garlic and onions. In Asian countries where diets are low in fat and high in soy and green tea, the incidence of breast, prostate, and colorectal cancer is low. Diets rich in fruits and vegetables lower the incidence of heart disease and of certain types of cancer. Studies comparing diets in various countries suggest that intake of fruits and vegetables may be more important than differences in dietary fat. Yet nutritionists estimate that in the United States only about 1 in 11 people eats the daily recommended 2 to 6.5 cups of fruits and vegetables a day (the equivalent of 4 to 13 servings). In the United States, the National Fruit and Vegetable Program (a public private partnership that includes government agencies, industry, and not-forprofit groups) promotes a public health initiative known as Fruit & Veggies–More Matters. This group is committed to increasing the number of daily servings of fruits and vegetables based on current dietary guidelines.

Review ■ ■ ■ ■

How do complex carbohydrates affect the body? What is the function of glucose? of essential amino acids? What is the function of high-density lipoproteins (HDLs)? What measures can you take to lower your risk for heart disease?

47.4 ENERGY METABOLISM ■ ■ LEARNING OBJECTIVES 8 Contrast basal metabolic rate with total metabolic rate; write the basic energy equation for maintaining body weight and describe the consequences of altering it in either direction.

9 Identify components of the regulatory pathways that regulate food intake and energy homeostasis.

10 In general terms, describe the effects of malnutrition, including both overnutrition and undernutrition.

The amount of energy liberated by the body per unit time is a measure of its metabolic rate. Much of the energy expended by the body is ultimately converted to heat. Metabolic rate may be expressed either in kilocalories of heat energy expended per day or as a percentage above or below a standard normal level. The basal metabolic rate (BMR) is the rate at which the body releases heat as a result of breaking down fuel molecules. BMR is the body’s basic cost of living, that is, the rate of energy use during resting conditions. This energy is required to maintain body functions such as heart contraction, breathing, and kidney function. A person’s total metabolic rate is the sum of his or her BMR and the energy used to carry on daily activities. A laborer has a greater total

metabolic rate than an account executive who is inactive most of the day and who does not exercise regularly. An average-sized man who does not exercise and who sits at a desk all day expends about 2100 kcal daily (women expend less energy). If the food the individual eats each day also contains about the same number of kilocalories, the body will be in a state of energy balance; that is, energy input will equal energy output. This is an extremely important concept because body weight remains constant when: energy input = energy output

When energy output is greater than energy input, stored fat is burned and body weight decreases. People gain weight when they take in more energy in food than they expend in daily activity, in other words, when: energy input > energy output

Energy metabolism is regulated by complex signaling The hypothalamus regulates energy metabolism and food intake. Biologists have identified several signaling molecules that form complex signaling pathways involving the hypothalamus and digestive system. For example, several appetite regulators signal us to eat or to stop eating. When the stomach is empty, it secretes the peptide hormone ghrelin (grell¿-in), which stimulates appetite. Ghrelin activates neurons in the hypothalamus that produce the appetite-stimulating neurotransmitter neuropeptide Y (NPY). The gastrointestinal tract releases several peptides that signal satiety and suppress food intake. Peptide YY, secreted by the intestine, signals the hypothalamus to stop secreting NPY. The decrease in NPY slows gastric emptying. Other compounds that signal satiety include the gastrointestinal hormone cholecystokinin (CCK) and melanocortins released by the hypothalamus. Leptin and insulin are also critical in regulating energy metabolism. Leptin, a hormone released by adipose tissue, is important in regulating normal body weight. Leptin signals centers in the brain about the status of energy stores in adipose tissue. The brain then adjusts feeding behavior and energy metabolism. The pancreatic hormone insulin is secreted when the concentration of glucose and other nutrients increases after we eat. Insulin stimulates cells, especially liver, muscle, and fat cells, to take in glucose from the blood and use or store it. The actions of insulin inhibit the hypothalamus from secreting NPY. Thus, insulin inhibits further food intake. The hypothalamus integrates input from these signaling molecules with other information, such as the concentration of nutrients circulating in the blood. The brain then sends signals that adjust feeding behavior and metabolism so that homeostasis of stored fat and fuel metabolism is maintained. When food is available and energy stores are adequate, energy use increases and fat stores are mobilized. Energy intake and glucose production in the liver are inhibited. In contrast, when food intake is low, as during dieting or starvation, the hypothalamus increases secretion of

underexercising lead to obesity, and psychosocial factors influence these behaviors. A sedentary lifestyle combined with the ready availability of high-energy foods contributes to the problem. One researcher cleverly described the roots of the obesity problem as the combination of “computer chips and potato chips.” For every 9.3 kcal of excess food taken into the body, about 1 g of fat is stored. (An excess of about 140 kcal, less than a typical candy bar, per day for a month results in a 1-pound weight gain.)

neuropeptide Y. Appetite increases and metabolism slows, actions that help restore energy homeostasis.

Obesity is a serious nutritional problem Overnutrition can cause malnutrition. Obesity, the excess accumulation of body fat, is a serious form of malnutrition that has become a problem of epidemic proportions in affluent societies. The World Health Organization considers obesity among the most important global health problems. Obesity contributes to about 300,000 deaths annually in the United States and is the second leading preventable cause of death (second only to smoking). It is a major risk factor for heart disease, diabetes mellitus, certain types of cancer (including breast and colon cancers), hypertension, osteoarthritis, and liver and gallbladder disease. In the well-known Framingham study of more than 2000 men, those who were 20% overweight had a significantly higher mortality rate from all causes. Body mass index (BMI), now used worldwide as a measure of body size, is an index of weight in relation to height. It is calculated by dividing the square of the weight (kg2) by height (m). The equivalent in the United States is 4.89 times the weight (lb) divided by the square of the height (ft2). An adult who has a BMI between 25 and 29.9 is considered overweight. An adult who has a BMI of 30 or more is considered obese. Approximately 33% of U.S. adults are overweight and another 34% of U.S. adults are obese. An estimated 17% of U.S. children and adolescents are overweight. Each of us appears to have a set point, or steady state, around which body weight is regulated. Dieting can be difficult because when BMI decreases below an individual’s set point, energy-conserving mechanisms are activated and energy expenditure decreases.

Undernutrition can cause serious health problems Millions of people do not have enough to eat or do not eat a balanced diet. Individuals suffering from undernutrition are malnourished. They may feel weak, are easily fatigued, and are highly susceptible to infection as a result of depressed immune function. Iron, calcium, and vitamin A are commonly deficient, but essential amino acids are the nutrients most often deficient in the diet. Millions of people suffer from poor health and lowered resistance to disease because of protein deficiency. Children’s physical and mental development are retarded when these essential building blocks of cells are not provided in the diet. Because these children’s bodies cannot manufacture antibodies (which are proteins) and cells needed to fight infection, common childhood diseases, such as measles, whooping cough, and chickenpox, are often fatal in children suffering from protein malnutrition.

© 1995 Amgen Inc.

Many factors affect weight In addition to an array of signaling molecules that affect energy metabolism and body weight, researchers have identified several gene mutations that affect food intake and energy homeostasis. For example, mice with a mutated allele at the ob locus become grossly obese. The normal ob gene product is leptin. When researchers inject leptin into grossly obese mice, the appetites of the mice decrease and their energy use increases. They lose weight, mainly from loss of body fat (FIG. 47-15). Unfortunately, most obese humans are resistant to the actions of leptin. The number of fat cells in the adult is apparently determined mainly by the amount of fat stored during infancy and childhood. When we are overfed early in life, abnormally large numbers of fat cells are formed. Later in life, these fat cells may be fully stocked with excess lipids or may shrink in size, but they are always there. People with such increased numbers of fat cells are thought to be at greater risk for obesity than are those with normal numbers. Eating activates the dopamine reward circuit in the brain. (Recall that this circuit is also activated in drug addiction.) The amount of dopamine released corresponds to the intensity of pleasure experienced by eating the food. One hypothesis holds that obese individuals may have an underactive dopamine reward system. They overeat to compensate and may develop a food addiction. Although many factors involved in obesity are inherited, behavior is critically important. High-calorie diets, overeating, and

Researchers have identified targets for obesity treatment As researchers discover the genetic and biochemical mechanisms that regulate energy metabolism, they provide potential targets for the development of pharmacological treatments for obesity. The following are among the many targets for drug action that are the focus of research: (1) reduce appetite so that food intake is decreased; (2) block absorption of fat; (3) uncouple metabolism from energy production so that food energy is dissipated as heat (see Inquiring About: Electron Transport and Heat, in Chapter 8); and (4) block receptors for molecules that send signals leading to increased energy storage.

FIGURE 47-15 Effect of leptin Mutant ob mice before (left) and after (right) treatment with leptin.

P. Pittet/United Nations Food and Agricultural Organization

In young children, severe protein malnutrition results in the condition known as kwashiorkor. This Ashanti (West African) word means “first-second.” It refers to the situation in which a first child is displaced from its mother’s breast when a younger sibling is born. The older child is then given a diet of starchy cereal or cassava that is deficient in protein. Growth becomes stunted, muscles are wasted, and edema develops (as displayed by a swollen belly); the child becomes apathetic and anemic, with an impaired metabolism (FIG. 47-16). Without essential amino acids, digestive enzymes cannot be manufactured, so what little protein is ingested cannot be digested. Deficiencies in trace elements and antioxidants have also been associated with the development of kwashiorkor.

Review ■

■ ■

Write an equation to describe energy balance and explain what happens when the equation is altered in either direction. What is body mass index (BMI)? What is the function of ghrelin? of leptin?

FIGURE 47-16 Protein deficiency Millions of children suffer from kwashiorkor, a disease caused by severe protein deficiency. Note the characteristic swollen belly, which results from fluid imbalance.

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S U M M A RY: F O C US O N L E A R N I N G O B J E C T I V E S

47.1 (page 1013) 1 Describe food processing, including ingestion, digestion, absorption, and egestion or elimination, and compare the digestive system of a cnidarian (such as Hydra) with that of an earthworm and vertebrate. ■ Nutrition is the process of taking in and using food. Feeding is the selection, acquisition, and ingestion of food, that is, taking food into the body. Digestion is the process of breaking down food mechanically and chemically. Nutrients pass through the lining of the digestive tract and into the blood by absorption. Food that is not digested and absorbed is discharged from the body by egestion or elimination. In cnidarians and flatworms, food is digested in the gastrovascular ■ cavity. This cavity has only one opening that serves as both mouth and anus. In more complex invertebrates and in all vertebrates, the digestive tract is a complete tube with an opening at each end. Digestion takes place as food passes through the tube. Various parts of the digestive tract are specialized to perform specific functions. Explore various digestive systems by clicking on the figure in CengageNOW.

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47.2 (page 1016) 2 Trace the pathway traveled by an ingested meal through the human digestive system and describe the structure and function of each organ involved. ■ Mechanical and enzymatic digestion of carbohydrates begins in the mouth. Mammalian teeth include incisors, which are adapted for biting and cutting food; canines adapted for tearing food; and

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premolars and molars adapted for crushing and grinding. Three pairs of salivary glands secrete saliva, which contains the enzyme salivary amylase that digests starch. As food is swallowed, it is propelled through the pharynx and esophagus. A bolus of food is moved along through the digestive tract by peristalsis, waves of muscular contraction that push food along. In the stomach, food is mechanically digested by vigorous churning, and proteins are enzymatically digested by the action of pepsin in the gastric juice. Rugae are folds in the stomach wall that expand as the stomach fills with food. Gastric glands secrete hydrochloric acid and pepsinogen, the precursor of pepsin. After several hours, a soup of partly digested food, called chyme, leaves the stomach through the pylorus and enters the small intestine in spurts. Most enzymatic digestion takes place in the duodenum, which produces several digestive enzymes and receives secretions from the liver and pancreas. The liver produces bile, which emulsifies fats. The pancreas releases enzymes that digest carbohydrate, lipid, and protein as well as RNA and DNA. Pancreatic amylase digests complex carbohydrates. Pancreatic lipase degrades fats, and trypsin and chymotrypsin digest polypeptides to dipeptides. The large intestine, which consists of the cecum, colon, rectum, and anus, is responsible for eliminating undigested wastes. It also incubates bacteria that produce vitamin K and certain B vitamins. Learn more about human digestion by clicking on the figures in Cengage NOW.

3 Trace the step-by-step digestion of carbohydrate, protein, and lipid. Nutrients in chyme are enzymatically digested as they move through the digestive tract. Polysaccharides are digested to the disaccharide maltose by salivary and pancreatic amylases. Maltase in the small intestine splits maltose into glucose, the main product of carbohydrate digestion. ■ Proteins are split by pepsin in the stomach and by proteolytic enzymes in the pancreatic juice. Dipeptidases split small peptides to amino acids. Lipids are emulsified by bile salts and then hydrolyzed by pancreatic ■ lipase. 4 Describe the structural adaptations that increase the surface area of the digestive tract. ■ The surface area of the small intestine is greatly expanded by folds in its wall; by the intestinal villi, elongated projections of the mucosa; and by microvilli, projections of the plasma membrane of the epithelial cells of the villi. ■

Villi

ols. Fatty acids are converted to molecules of acetyl coenzyme A, which enter the citric acid cycle. Excess fatty acids are converted to triacylglycerol and stored as fat. ■ Lipids are transported as large molecular complexes called lipoproteins. Low-density lipoproteins (LDLs) deliver cholesterol to the cells. High-density lipoproteins (HDLs) collect excess cholesterol and transport it to the liver. ■ Proteins serve as enzymes and are essential structural components of cells. The best distribution of essential amino acids is found in the complete proteins of animal foods. Excess amino acids are deaminated by liver cells. Amino groups are converted to urea and excreted in urine; the remaining keto acids are converted to carbohydrate and used as fuel or converted to lipid and stored in fat cells. 7 Describe the nutritional functions of vitamins, minerals, and phytochemicals. ■ Vitamins are organic compounds required in small amounts for many biochemical processes. Many serve as components of coenzymes. Fat-soluble vitamins include vitamins A, D, E, and K. Watersoluble vitamins are the B and C vitamins. ■ Minerals are inorganic nutrients ingested as salts dissolved in food and water. Trace elements are minerals required in amounts less than 100 mg per day. ■ Phytochemicals are plant compounds that promote health. Many phytochemicals are antioxidants that destroy oxidants. which are reactive molecules, such as free radicals. These molecules damage DNA, proteins, and unsaturated fatty acids by snatching electrons.

47.4 (page 1029) 8 Contrast basal metabolic rate with total metabolic rate; write the basic

5 Compare lipid absorption with absorption of other nutrients. ■

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Nutrients are absorbed through the thin walls of intestinal villi. The hepatic portal vein transports amino acids and glucose to the liver. Fatty acids and monoacylglycerols enter epithelial cells in the intestinal lining, where they are reassembled into triacylglycerols. They are packaged into chylomicrons, droplets that also contain cholesterol and phospholipids and are covered with a protein coat. The lymphatic system transports chylomicrons to the blood circulation.

energy equation for maintaining body weight and describe the consequences of altering it in either direction. ■ Basal metabolic rate (BMR) is the body’s cost of metabolic living. Total metabolic rate is the BMR plus the energy used to carry on daily activities. ■ When energy (kilocalories) input equals energy output, body weight remains constant. When energy output is greater than energy input, body weight decreases. When energy input exceeds energy output, body weight increases. 9 Identify components of the regulatory pathways that regulate food intake and energy homeostasis. ■ Complex pathways of signaling molecules regulate food intake and energy metabolism. ■ Ghrelin and neuropeptide Y (NPY) stimulate appetite. Peptide YY, cholecystokinin (CCK), and melanocortins signal satiety. ■ The hormone leptin signals the brain about the status of energy stores in adipose tissue. Leptin and insulin inhibit food intake. Learn more about leptins and body weight by clicking on the figure in CengageNOW.

47.3 (page 1024) 6 Summarize the nutritional requirements for dietary carbohydrates, lipids, and proteins, and trace the fate of glucose, lipids, and amino acids after their absorption. Most carbohydrates are ingested in the form of polysaccharides— ■ starch and cellulose. Polysaccharides are referred to as complex carbohydrates. Fiber is mainly a mixture of cellulose and other indigestible carbohydrates. Carbohydrates are used mainly as an energy source. Glucose concentration in the blood is carefully regulated. Excess glucose is stored as glycogen and can also be converted to fat. ■ Lipids are used to provide energy, to form components of cell membranes, and to synthesize steroid hormones and other lipid substances. Most lipids are ingested in the form of triacylglycer-

10 In general terms, describe the effects of malnutrition, including both overnutrition and undernutrition. In obesity an excess amount of fat accumulates in adipose tissues. Obesity is a major risk factor for several major disorders, including heart disease and diabetes mellitus. A person gains weight by taking in more energy, in the form of kilocalories, than is expended in activity. Genes, signaling molecules that affect energy metabolism, and certain behaviors contribute to obesity. ■ Millions of people suffer from undernutrition, a form of malnutrition that causes fatigue and depressed immune function. Essential amino acids are the nutrients most often deficient in the diet. ■

T E S T YO U R U N D E R S TA N D I N G 1. Teeth adapted for crushing and grinding are (a) incisors (b) canines (c) premolars and molars (d) incisors and premolars (e) canines and molars

11. A hormone that stimulates gastric glands to secrete pepsinogen is (a) secretin (b) GIP (c) gastrin (d) cholecystokinin (CCK) (e) substance P

2. The layer of tissue that lines the lumen of the digestive tract is the (a) muscle layer (b) visceral peritoneum (c) parietal peritoneum (d) mucosa (e) submucosa

12. Label the diagram. Use Figure 47-4 to check your answers.

3. Which of the following are accessory digestive glands? (a) salivary glands (b) pancreas (c) liver (d) a, b, and c (e) a and b 4. Arrange the following into the correct sequence. 1. stomach 2. esophagus 3. pharynx 4. small intestine 5. colon (a) 2, 3, 1, 4, 5 (b) 3, 2, 1, 5, 4 (c) 3, 2, 5, 1, 4 (d) 2, 3, 1, 5, 4 (e) 3, 2, 1, 4, 5 5. Amylase is produced by the (a) pancreas and salivary glands (b) stomach and pancreas (c) colon and salivary glands (d) liver and pancreas (e) duodenum and pancreas 6. Which sequence most accurately describes the digestion of protein? 1. dipeptide 2. amino acid 3. protein 4. polypeptide 5. glycerol (a) 4, 3, 1, 2 (b) 3, 4, 1, 2 (c) 3, 4, 1, 5 (d) 4, 3, 5, 1, 2 (e) 2, 1, 4, 5 7. The surface area of the small intestine is increased by (a) folds in its wall (b) villi (c) microvilli (d) a, b, and c (e) a and b 8. Lipids are transported from the intestine to the liver by (a) chylomicrons (b) HDLs (c) LDLs (d) glycerol transporters (e) leptin 9. Most vitamins are (a) inorganic compounds (b) components of coenzymes (c) used as fuel (d) electrolytes (e) required by herbivores and carnivores but not omnivores 10. When energy input is greater than energy output, (a) weight loss occurs (b) weight remains stable (c) weight gain occurs (d) leptin secretion is inhibited (e) the ob gene is deactivated

CRITICAL THINKING 1. Design an experiment to test the hypothesis that the B vitamin pyridoxine is an essential nutrient in mice. 2. Proteolytic enzymes are produced in an inactive form. What, if any, benefit might this have? 3. EVOLUTION LINK. If you were presented with an unfamiliar animal and asked to determine its nutritional lifestyle, how would you do so? (Hint: Consider adaptations.)

4. SCIENCE, TECHNOLOGY, AND SOCIETY. Investigators are unraveling the biochemical pathways that help regulate body weight. How might their work lead to a cure for obesity? How might a cure for obesity affect our society? Additional questions are available in CengageNOW at www.cengage.com/ login.

48

Osmoregulation and Disposal of Metabolic Wastes

McMurray Photography

Most terrestrial animals inhabit areas near water sources. Zebra, wildebeest, and a variety of birds drink at a lake in Ngorongoro Crater in Tanzania.

KEY CONCEPTS

W

48.1 Osmoregulation is the process by which organisms

ganisms on our planet. Like the animals in the photograph, most animals

control the concentration of water and salt in the body so that their body fluids are maintained within homeostatic limits.

need a dependable source of water with which to replenish their body

48.2 During metabolism, most animals produce water, carbon dioxide, and nitrogenous wastes, including ammonia, uric acid, and urea.

ater, the most abundant molecule both in the cell and on Earth’s surface, shapes life and is a critical factor in the distribution of or-

fluids. For this reason, animals often inhabit areas near water sources. Ions are dissolved in water, the medium in which most metabolic reactions take place. The volume and ionic composition of an animal’s body fluids must be maintained within homeostatic limits. Through natural se-

48.3 Many invertebrates have nephridial organs that

lection a variety of homeostatic mechanisms have evolved that regulate

carry on osmoregulation and excretion; insects and spiders have Malpighian tubules that excrete metabolic wastes and conserve water.

the volume and composition of fluids in the internal environment.

48.4 Freshwater, marine, and terrestrial animals have different adaptations for osmoregulation that meet the challenges of these diverse environments.

Many small animals live in the ocean and get their food and oxygen directly from the sea water that surrounds them; they release waste products into the sea water. In larger animals and in most terrestrial animals, the cells are bathed by fluid that serves as an internal sea. Terrestrial ani-

48.5 The vertebrate kidney maintains water and electro-

mals have a continuous need to conserve water. Water loss from the body

lyte balance and excretes metabolic wastes; the nephron is the functional unit of the vertebrate kidney. Several hormones interact to regulate kidney function.

must be carefully regulated, and water lost must be replaced. Water is taken in with food and drink and is also produced in metabolic reactions.

Two processes that help maintain fluid and electrolyte (salt) homeostasis in animals are osmoregulation and excretion, the disposal of metabolic wastes. In this chapter we discuss these processes in both invertebrates and vertebrates inhabiting various environments. Then we focus on the structure and functions of the mammalian kidney. We end the chapter with a description of the hormones that regulate kidney function.

48.1 MAINTAINING FLUID AND ELECTROLYTE BALANCE ■ ■ LEARNING OBJECTIVE 1

Describe how the processes of osmoregulation and excretion contribute to fluid and electrolyte homeostasis.

Intracellular fluid, the fluid within cells, accounts for most of the body fluid. Extracellular fluid, the fluid outside the cells, includes interstitial fluid (the fluid between cells), lymph, and blood plasma (or hemolymph). In vertebrates, blood plasma, which is mainly water, transports nutrients, gases, waste products, and other materials throughout the body. Interstitial fluid forms from the blood plasma and bathes all the cells. Excess water evaporates from the body surface or is excreted by specialized structures. Electrolytes are compounds such as inorganic salts, acids, and bases that form ions in solution. Electrolytes are very important solutes in body fluids. Many homeostatic mechanisms regulate fluid and electrolyte balance. Recall from Chapter 5 that osmosis is the diffusion of water through a selectively permeable membrane. The net flow of water is from a more dilute solution (one with a lower solute concentration) to a less dilute solution (one with a higher solute concentration). If the solute concentrations of two solutions are equal, they are isotonic to each other. If solution A has a greater solute concentration than solution B, solution A is hypertonic to solution B. Solution B, with a lower solute concentration relative to solution A, is hypotonic to solution A. Remember that the total solute concentration of a solution is responsible for the direction of water movement. The osmotic pressure of a solution is the pressure that must be exerted on the hypertonic side of a selectively permeable membrane to prevent net movement of water from the hypotonic side. An osmole is related to a mole (see Chapter 2), but takes into account the number of particles produced when a solute dissolves. For example, glucose dissolves to give only one kind of particle, so a mole of glucose in solution is one osmole. A mole of NaCl in solution produces two kinds of particles (Na+ and Cl−), so a mole of NaCl is two osmoles. Both types of ions affect the osmotic pressure of the solution. A milliosmole is 1/1000 of an osmole. Osmolarity is a measure of the number of osmoles of solute per liter of solution. Solutions with the same osmolarity are described as isoosmotic. If solution X has a higher osmolarity than solution Y, it is described as hyperosmotic. Solution Y would be described as hypoosmotic relative to solution X.

Osmoregulation is the process by which organisms control the concentration of water and salt so that their body fluids do not become too dilute or too concentrated. Excretion is the process of ridding the body of metabolic wastes. Excretory systems have evolved that function in both osmoregulation and in disposal of metabolic wastes. Excretory systems rid the body of excess water and ions, metabolic wastes, and harmful substances. As we will discuss, hormones are important signaling molecules in these regulatory processes.

Review ■ ■

What is osmoregulation? What is excretion?

48.2 METABOLIC WASTE PRODUCTS ■ ■ LEARNING OBJECTIVE 2 Contrast the benefits and costs of excreting ammonia, uric acid, or urea.

Metabolic wastes must be excreted so that they do not accumulate and reach concentrations that would disrupt homeostasis. The principal metabolic waste products produced by most animals are water, carbon dioxide, and nitrogenous wastes, those that contain nitrogen. Carbon dioxide is excreted mainly by respiratory structures (see Chapter 46). In terrestrial animals, some water is also lost from respiratory surfaces. However, specialized excretory organs, such as kidneys, remove and excrete most of the water and nitrogenous wastes. Nitrogenous wastes include ammonia, uric acid, and urea. Recall that amino acids and nucleic acids contain nitrogen. During the metabolism of amino acids, the nitrogen-containing amino group is removed (in a process known as deamination) and converted to ammonia (FIG. 48-1). However, ammonia is highly toxic. Some aquatic animals excrete it into the surrounding water before it can build up to toxic concentrations in their tissues. A few terrestrial animals, including some snails and wood lice, vent it directly into the air. But many animals, humans included, convert ammonia to some less toxic nitrogenous waste such as uric acid or urea. Uric acid is produced both from ammonia and by the breakdown of nucleotides from nucleic acids. Uric acid is insoluble in water and forms crystals that are excreted as a crystalline paste, so little fluid loss results. This is an important water-conserving adaptation in many terrestrial animals, including insects, certain reptiles, and birds. Also, because uric acid is not toxic and can be safely stored, its excretion is an adaptive advantage for species whose young begin their development enclosed in eggs. Urea, the principal nitrogenous waste product of amphibians and mammals, is synthesized in the liver from ammonia and carbon dioxide by a sequence of reactions known as the urea cycle. Like the formation of uric acid, these reactions require specific enzymes and the input of energy by the cells. Compared with the

Amino acids NH2 H

O

C

C

48.3 OSMOREGULATION AND EXCRETION IN INVERTEBRATES

Nucleic acids

OH

R

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Deamination

3 Compare osmoconformers and osmoregulators. 4 Describe protonephridia, metanephridia, and Malpighian tubules, and compare their functions. Keto acids O

Ammonia NH3

C

C

R Urea cycle

Ammonia NH3

Purines O OH

15 steps

H2N

Urea NH2 C O

HN O

C

Uric acid O H C N C N H

C

C

O

N H

More energy needed to produce More water needed to excrete

FIGURE 48-1 Formation of nitrogenous wastes Deamination of amino acids and metabolism of nucleic acids produce nitrogenous wastes. Ammonia is the first metabolic product of deamination. Many aquatic animals excrete ammonia, but terrestrial animals convert it to urea and uric acid, which are less toxic. Most amphibians and mammals convert ammonia to urea via the urea cycle. Insects, many reptiles, and birds convert ammonia to uric acid. Energy is required to convert ammonia to urea and uric acid, but less water is required to excrete these wastes.

energy cost of producing ammonia, that of producing urea and uric acid is high. Urea has the advantage of being far less toxic than ammonia and can accumulate in higher concentrations without causing tissue damage; thus, it can be excreted in more concentrated form. Because urea is highly soluble, however, it is dissolved in water, and more water is needed to excrete urea than to excrete uric acid.

Review ■

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What are the principal types of nitrogenous wastes? Give examples of animals that excrete each type. What are the benefits of excreting nitrogenous wastes in the form of uric acid? in the form of urea?

The ocean is a stable environment, and its salt concentration does not vary much. The electrolyte concentration in the cells and body fluids of marine sponges and cnidarians is very similar to sea water, so these animals do not require specialized excretory structures. They expend little or no energy in excreting wastes because wastes simply diffuse from their cells to the external environment and are washed away by water currents. When water stagnates and currents do not wash away wastes, aquatic environments, such as coral reefs, are damaged by the waste accumulation. The body fluids of most marine invertebrates are in osmotic equilibrium with the surrounding sea water. These animals are known as osmoconformers because the concentration of their body fluids varies along with changes in the sea water. However, many osmoconforming marine animals still regulate some ions in their body fluids. Coastal habitats, such as estuaries that contain brackish water, are much less stable environments than is the open ocean. Salt concentrations change frequently with shifting tides. Many invertebrates and vertebrates that inhabit these environments are osmoregulators. These animals have homeostatic mechanisms that maintain an optimal salt concentration in their tissues regardless of changes in the salt concentration of their surroundings. In a coastal environment where fresh water enters the ocean, the water may have a lower salt concentration than an animal’s body fluids. Water osmotically moves into the animal’s body, and salt diffuses out. An animal adapted to this environment has excretory structures that remove the excess water. Certain polychaete worms and the blue crab are among the animals that can be osmoconformers or osmoregulators depending on environmental conditions. Dehydration is a constant threat to terrestrial animals. Because their fluid concentration is higher than that of the air around them, they tend to lose water by evaporation from both the body surface and respiratory surfaces. Terrestrial animals may also lose water as wastes are excreted. As animals moved onto the land, natural selection favored the evolution of structures and processes that conserve water. Excretory systems help maintain fluid and electrolyte homeostasis by selectively adjusting the concentrations of salts and other substances in blood and other body fluids. Typically, an excretory system collects fluid, generally from the blood or interstitial fluid. It then adjusts the composition of this fluid by selectively returning needed substances to the body fluid. Finally, the body releases the adjusted excretory product containing excess or potentially toxic substances.

Nucleus

Flame cells

Cytoplasm Cilia (“flame”) Protonephridial network

Movement of interstitial fluid

Nephridiopores

Excretory tubule

Excretory tubule

(b) Interstitial fluid enters flame cells and passes through a series of tubules. Excess fluid leaves the body through nephridiopores.

(c) A single flame cell.

(a) The protonephridia of a planarian function mainly in osmoregulation. These organs form a system of branching tubules.

FIGURE 48-2 Protonephridia of a flatworm

Nephridial organs are specialized for osmoregulation and/or excretion Nephridial organs, or nephridia, are structures that function in osmoregulation and excretion in many invertebrates, including flatworms, nemerteans, rotifers, annelids, mollusks, and lancelets. Each nephridial organ consists of simple or branching tubes that typically open to the outside of the body through pores, called nephridiopores. Two types of nephridial organs are protonephridia and metanephridia. In flatworms and nemerteans, metabolic wastes pass through the body surface by diffusion, but these animals also have protonephridia (FIG. 48-2). The protonephridia are composed of tubules with no internal openings. Their enlarged blind ends consist of flame cells with brushes of cilia, so named because their constant motion reminded early biologists of flickering flames. The flame cells lie in the interstitial fluid that bathes the body cells. Fluid enters the flame cells, and the beating of the cilia propels the fluid through the tubules. Excess fluid leaves the body through nephridiopores. Most annelids and mollusks have more complex nephridial organs called metanephridia (FIG. 48-3). Each segment of an earthworm has a pair of metanephridia. A metanephridium is a tubule open at both ends. The inner end opens into the coelom as a ciliated funnel, and the outer end opens to the outside through a nephridiopore. A network of capillaries surrounds each tubule.

Tubule Posterior

Anterior

Gut Funnel

Septum

Nephridiopore

Capillary network

FIGURE 48-3 Metanephridium of an earthworm Each metanephridium consists of a ciliated funnel opening into the coelom, a coiled tubule, and a nephridiopore opening to the outside. This 3-D internal view shows parts of three segments of the earthworm body.

Fluid from the coelom passes into the tubule, bringing with it whatever it contains—glucose, salts, or wastes. As fluid moves through the tubule, needed materials (such as water and glucose) are removed from the fluid by the tubule and are reabsorbed by the capillaries, leaving the wastes behind. In this way, urine that contains concentrated wastes is produced.

Gut

Malpighian tubules

Waste

Malpighian tubules conserve water The excretory system of insects and spiders consists of several hundred Malpighian tubules (FIG. 48-4). Malpighian tubules are slender extensions of the gut wall. Their blind ends lie in the hemocoel (blood cavity) and are bathed in hemolymph. Cells of the tubule wall actively transport uric acid, potassium ions, and some other substances from the hemolymph into the tubule lumen. Other solutes and water follow by diffusion. The Malpighian tubules empty into the gut. Water, some salts, and other solutes are reabsorbed into the hemolymph by a specialized epithelium in the rectum. Uric acid, the major waste product, is excreted as a semidry paste with a minimum of water loss. Because Malpighian tubules effectively conserve body fluids, they have contributed to the success of insects in terrestrial environments.

Review ■

■ ■ LEARNING OBJECTIVES 6

Hindgut

Rectum

Water and needed ions

FIGURE 48-4 Malpighian tubules of an insect The slender Malpighian tubules have blind ends that extend into the hemocoel. Their cells transfer uric acid and some ions from the hemolymph to the cavity of the tubule. Water follows by diffusion. The wastes are discharged into the gut. The epithelium that lines the rectum (part of the hindgut) actively reabsorbs most of the water and needed ions.

How are nephridial organs and Malpighian tubules alike? How are they different?

48.4 OSMOREGULATION AND EXCRETION IN VERTEBRATES 5

Midgut

Relate the function of the vertebrate kidney to the success of vertebrates in a wide variety of habitats. Compare adaptations for osmoregulation in freshwater fishes, marine bony fishes, sharks, marine mammals, and terrestrial vertebrates.

Vertebrates live successfully in a wide range of habitats—in fresh water, the ocean, tidal regions, and on land, even in extreme environments such as deserts. In response to the requirements of these diverse environments, adaptations have evolved for regulating salt and water content and for excreting wastes. The main osmoregulatory and excretory organ in most vertebrates is the kidney. The kidney excretes most nitrogenous wastes and helps maintain fluid balance by adjusting the salt and water content of the urine. The skin, lungs or gills, and digestive system also help maintain fluid balance and dispose of metabolic wastes.

Freshwater vertebrates must rid themselves of excess water As fishes began to move into freshwater habitats about 470 million years ago (mya), there must have been strong selection for ad-

aptations that promoted effective osmoregulation. The evolution of body fluids more dilute than sea water was a major adaptation. However, the salt concentration of the body fluids of these fishes is still higher than that of the fresh water that surrounds them. They are hypertonic to their watery environment. As a result, water moves into the body, and they are in constant danger of becoming waterlogged. Freshwater fishes are covered by scales and a mucous secretion that retard the passage of water into the body. However, water constantly enters through the gills and through the mouth (with food). How do these animals meet this challenge? Some water leaves the body through the gills. Furthermore, the kidneys of freshwater fishes are adapted to excrete large amounts of dilute urine (FIG. 48-5a). The kidneys have large glomeruli, capillary clusters that filter the blood and produce urine. Water entry is only part of the challenge of osmoregulation in freshwater fishes. These animals also tend to lose salts by diffusion through the gills into the surrounding water. To compensate, special gill cells have evolved that actively transport salts (mainly sodium chloride) from the water into the body. The gills excrete most of the nitrogenous wastes of freshwater fishes, but the kidneys are also important excretory organs. Ammonia is the main nitrogenous waste; about 10% of nitrogenous wastes are excreted as urea. Most amphibians are at least semi-aquatic, and their mechanisms of osmoregulation are similar to those of freshwater fishes. They, too, produce large amounts of dilute urine. In its urine and through its skin, a frog can lose an amount of water equivalent to one-third of its body weight in one day. Active transport of salt into

the body by special cells in the skin compensates for loss of salt through skin and in urine.

Marine vertebrates must replace lost fluid Freshwater fishes adapted very successfully to their aquatic habitats. Thus, when some freshwater fishes returned to the sea about 200 mya, their blood and body fluids were less salty than their surroundings; that is, they were hypotonic to their environment. These fishes lose water osmotically and take in salt. To compensate for fluid loss, many marine bony fishes drink sea water (FIG. 48-5b). They retain the water and excrete salt by the action of specialized cells in their gills. The gills are also responsible for ammonia excretion. Very little urine is excreted by the kidneys, which have only small (or no) glomeruli. Marine cartilaginous fishes (sharks and rays) have different osmoregulatory adaptations that allow them to tolerate the salt concentrations of their environment. These animals accumulate and tolerate urea (FIG. 48-5c). Their tissues are adapted to function at concentrations of urea that would be toxic to most other animals. The high urea concentration makes the body fluids slightly hypertonic to sea water, resulting in a net inflow of water. Marine cartilaginous fishes have well-developed kidneys that excrete a large volume of urine. Excess salt is excreted by the kidneys and, in many species, by a rectal gland. Certain reptiles and marine birds ingest sea water and take in a lot of salt in their food. Glands in their heads excrete salt. These salt glands are usually inactive; they function only in response to osmotic stress. Whales, dolphins, and other marine mammals ingest sea water along with their food. Their kidneys produce concentrated urine, much saltier than sea water. This is an important physiological adaptation, especially for marine carnivores. The high-protein diet of these animals results in the production of large amounts of urea, which must be excreted in urine without losing much water.

Terrestrial vertebrates must conserve water Adult amphibians generally inhabit moist environments. They excrete urea and reabsorb some water from the urinary bladder. Amniotes (reptiles, birds, and mammals) have more effective adaptations for life on land. Their skin minimizes water loss by evaporation, and many amniotes excrete uric acid, which requires very little water. Because birds and mammals are endothermic (maintain a constant body temperature) and have a high rate of metabolism, they produce a relatively large volume of nitrogenous wastes. They have numerous adaptations, including very efficient kidneys, for conserving water. An extreme example is the desert-dwelling kangaroo rat, which obtains most of its water from its own metabolism. (Recall from Chapter 8 that aerobic respiration produces water. The kangaroo rat’s kidneys are so efficient that it loses little fluid as urine.) Birds conserve water by excreting nitrogen as uric acid and by efficiently reabsorbing water from the cloaca and intestine. Mammals excrete urea. Their kidneys produce very concentrated (hypertonic) urine.

KEY POINT

Fishes have a variety of adaptations for regulating water and salt concentration. Loses salt by diffusion

Water gain by osmosis

Drinks no water Salt uptake by gills

Kidney with large glomeruli

Large volume of hypotonic urine

(a) Freshwater fishes live in a hypotonic medium. Water continuously enters the body, and salts diffuse out. These fishes excrete large quantities of dilute urine and actively transport salts in through the gills. Gains salts by diffusion

Water loss by osmosis

Drinks salt water Salt excreted through gills

Small volume of isotonic urine Kidney with small or no glomeruli

(b) Marine fishes live in a hypertonic medium. They lose water by osmosis. They gain salts from the sea water they drink and by diffusion. To compensate, the fish drinks water, excretes the salt, and produces a small volume of urine. Water gain by osmosis Salt-excreting gland

Salts diffuse in through gills

Some salt water swallowed with food

Kidney with large glomeruli— reabsorbs urea

Large volume of hypotonic urine

(c) Cartilaginous fishes (sharks and rays). The shark kidney reabsorbs urea in high enough concentration that its tissues become hypertonic to the surrounding medium. As a result, water enters the shark by osmosis. The shark excretes a large quantity of dilute urine.

FIGURE 48-5 Osmoregulation in fishes

In terrestrial vertebrates, the lungs, skin, and digestive system are important in osmoregulation and waste disposal (FIG. 48-6). Most carbon dioxide is excreted by the lungs. In birds and mammals, some water is lost from the body as water vapor in exhaled air. Although primarily concerned with the regulation of body

Some water loss by evaporation

Blood filtered; water reabsorbed by kidney

Review ■

■

Drinks water

What type of osmoregulatory challenge is faced by marine fishes? by freshwater fishes? What mechanisms have evolved to meet these challenges? What adaptations in aquatic birds and mammals help meet the challenges of osmoregulation and metabolic waste disposal? in terrestrial birds and mammals?

48.5 THE URINARY SYSTEM ■ ■ LEARNING OBJECTIVES

Hypertonic urine

7 Describe (or label on a diagram) the organs of the mammalian urinary system and give the functions of each.

(a) The kidney of terrestrial vertebrates conserves water by reabsorbing it. Birds and mammals can produce concentrated urine.

8 Describe (or label on a diagram) the structures of a nephron (including associated blood vessels) and give the functions of each structure.

9 Trace a drop of filtrate from Bowman’s capsule to its release from the body as urine.

10 Describe the hormonal regulation of fluid and electrolyte balance Wastes produced

LIVER

ALL CELLS Hemoglobin breakdown

Breakdown of nucleic acids Deamination of amino acids Uric acid Wastes

by antidiuretic hormone (ADH), the renin–angiotensin–aldosterone system, and atrial natriuretic peptide (ANP).

Urea

Cellular respiration

Bile pigments

Water

Carbon dioxide

Organs of excretion KIDNEY

Excretion

Urine

DIGESTIVE SYSTEM Feces

SKIN

Sweat

LUNGS Exhaled air containing water vapor and carbon dioxide

(b) Disposal of metabolic wastes in humans and other terrestrial mammals. To conserve water, mammals produce a small amount of hypertonic urine. Nitrogenous wastes are produced by the liver and transported to the kidneys. All cells produce carbon dioxide and some water during cellular respiration. These waste products are excreted by the lungs.

FIGURE 48-6 Excretory organs in terrestrial vertebrates

temperature, the sweat glands of humans and some other mammals excrete 5% to 10% of all metabolic wastes. The liver produces both urea and uric acid, which are transported by the blood to the kidneys. Most of the bile pigments produced by the breakdown of red blood cells are excreted by the liver into the intestine. From the intestine they pass out of the body with the feces.

The mammalian urinary system consists of the kidneys, the urinary bladder, and associated ducts. The overall structure of the human urinary system is shown in FIGURE 48-7. Located just below the diaphragm in the “small of the back,” the kidneys look like a pair of giant, dark red lima beans, each about the size of a fist. Each kidney is covered by a connective tissue capsule (FIG. 48-8). The outer portion of the kidney is the renal cortex; the inner portion is the renal medulla. The renal medulla contains eight to ten coneshaped structures called renal pyramids. The tip of each pyramid is a renal papilla. Each papilla has several pores, the openings of collecting ducts. As urine is produced, it flows from collecting ducts through a renal papilla and into the renal pelvis, a funnel-shaped chamber. Urine then flows into one of the paired ureters, ducts that connect each kidney with the urinary bladder. The urinary bladder is a remarkable organ capable of holding (with practice) up to 800 mL (about a pint and a half) of urine. Emptying the bladder changes it from the size of a small melon to that of a pecan. This feat is made possible by the smooth muscle and specialized epithelium of the bladder wall, which is capable of great shrinkage and stretching. During urination, urine is released from the bladder and flows through the urethra, a duct leading to the outside of the body. In the male, the urethra is lengthy and passes through the penis. Semen, as well as urine, passes through the male urethra. In the female, the urethra is short and transports only urine. Its opening to the outside is just above the opening of the vagina. The length of the male urethra discourages bacterial invasions of the bladder. This length difference helps explain why bladder infections are more common in females than in males. In summary, urine flows through the following structures: kidney (through renal pelvis) ¡ ureter ¡ urinary bladder ¡ urethra

▲

FIGURE 48-7 Animated The human urinary system

The kidneys produce urine, which passes through the ureters to the urinary bladder for temporary storage. During urination, the urethra conducts urine from the bladder to the outside of the body through an opening, the external urethral orifice.

Adrenal gland Left renal artery

Right kidney Right renal vein

Left kidney Inferior vena cava

Abdominal aorta

Ureteral orifices Right and left ureters Urinary bladder Urethra

External urethral orifice

Juxtamedullary nephron

Distal convoluted tubule

Cortical nephron

Renal pyramids (medulla) Capsule Capsule Renal cortex Renal medulla

Proximal convoluted tubule Renal cortex

Renal artery

Glomerulus Bowman’s capsule

Artery and vein Loop of Henle

Renal vein Renal pelvis

Renal medulla

Ureter

(a) Internal structure of the kidney. The outer region of the kidney is the cortex; the inner region is the medulla. As urine is produced, it flows into the renal pelvis and leaves the kidney through the ureter.

FIGURE 48-8 Animated Structure of the kidney

Collecting duct Papilla

(b) Juxtamedullary and cortical nephrons. Longitudinal section showing the location and structure of the two main types of nephrons.

a glomerulus. Three main regions of the renal tubule are (1) the proximal convoluted tubule (or simply, proximal tubule), which conducts the filtrate (the fluid filtered from the blood) from Bowman’s capsule; (2) the loop of Henle, an elongated, hairpin-shaped portion; and (3) the distal convoluted tubule (or distal tubule), which conducts the filtrate to a collecting duct. Thus, filtrate passes through the following structures:

The nephron is the functional unit of the kidney The principal function of the kidneys is to help maintain homeostasis by regulating fluid balance and excreting metabolic wastes. However, the kidneys have other functions as well. They produce the enzyme renin, which helps regulate fluid balance and blood pressure (discussed later in the chapter). They also produce at least two hormones: erythropoietin, which stimulates red blood cell production, and 1,25-dihydroxy-vitamin D3 , which stimulates calcium absorption by the intestine. Each kidney has up to one million functional units called nephrons. A nephron consists of a cuplike Bowman’s capsule connected to a long, partially coiled renal tubule (FIG. 48-9). Positioned within Bowman’s capsule is a cluster of capillaries called

Bowman's capsule

Bowman’s capsule ¡ proximal convoluted tubule ¡ loop of Henle ¡ distal convoluted tubule ¡ collecting duct

The human kidney has two types of nephrons: the more numerous (85%) cortical nephrons and the more internal juxtamedullary nephrons (see Fig. 48-8b). Cortical nephrons have

Distal tubule

Proximal tubule

Bowman's capsule

Proximal tubule

Glomerulus Efferent arteriole Afferent arteriole Podocyte

Peritubular capillaries Afferent arteriole

Distal tubule

Collecting duct From renal artery

To renal vein

Loop of Henle

Juxtaglomerular apparatus Efferent arteriole

(b) Cutaway view of Bowman’s capsule. Note that the distal tubule is adjacent to the afferent and efferent arterioles. The juxtaglomerular apparatus (discussed later in the chapter) is a small group of cells located in the walls of the tubule and arterioles.

100 μm

(c) Low-power SEM of a portion of the kidney cortex. The tissue was treated to remove some structures and to show glomeruli and associated blood vessels.

Afferent arteriole

Distal tubule

Steve Gschmeissner/Photo Researchers, Inc.

(a) Location and the basic structure of a nephron. Urine forms by filtration of the blood in the glomerulus and by adjustment of the filtrate as it passes through the series of tubules that drain Bowman’s capsule.

To renal pelvis

Glomerular capillaries

FIGURE 48-9 Nephron structure

terials returned to the blood by the renal tubule. Blood from the peritubular capillaries enters small veins that eventually lead to the renal vein. In summary, blood circulates through the kidney in the following sequence:

relatively small glomeruli and are located almost entirely within the cortex or outer medulla. Juxtamedullary nephrons have large glomeruli, and their very long loops of Henle extend deep into the medulla. The loop of Henle consists of a descending limb that receives filtrate from the proximal convoluted tubule and an ascending limb, through which the filtrate passes on its way to the distal convoluted tubule. The juxtamedullary nephrons contribute to the ability of the mammalian kidney to concentrate urine. Excretion of urine that is hypertonic to body fluids is an important mechanism for conserving water. Blood is delivered to the kidney by the renal artery. Small branches of the renal artery give rise to afferent arterioles (afferent means “to carry toward”). An afferent arteriole conducts blood into the capillaries that make up each glomerulus. As blood flows through the glomerulus, some of the plasma is forced into Bowman’s capsule. You may recall that in a typical circulatory route, capillaries deliver blood into veins. Circulation in the kidneys is an exception because blood flowing from the glomerular capillaries next passes into an efferent arteriole. Each efferent arteriole conducts blood away from a glomerulus. The efferent arteriole delivers blood to a second capillary network, the peritubular capillaries surrounding the renal tubule. Blood is filtered as it flows through the first set of capillaries, those of the glomerulus. The peritubular capillaries receive ma-

KEY POINT

renal artery ¡ an afferent arteriole ¡ capillaries of a glomerulus ¡ an efferent arteriole ¡ peritubular capillaries ¡ small veins ¡ renal vein

Urine is produced by filtration, reabsorption, and secretion Urine, the watery discharge of the urinary system, is produced by a combination of three processes: filtration, reabsorption, and tubular secretion (FIG. 48-10). Filtration is not selective with regard to ions and small molecules Blood flows through the glomerular capillaries under high pressure, forcing more than 10% of the plasma out of the capillaries and into Bowman’s capsule. Filtration is similar to the mechanism whereby interstitial fluid is formed as blood flows through other capillary networks in the body. However, blood flow through

The glomerulus filters blood. As the filtrate moves through the renal tubule, its composition is adjusted by selective reabsorption and secretion. The adjusted filtrate is urine. REABSORPTION AND SECRETION Proximal tubule FILTRATION Bowman's capsule Glomerulus

REABSORPTION AND SECRETION

REABSORPTION OF H2O; URINE CONCENTRATED

Distal tubule

Collecting duct

Capillaries

Renal artery Renal vein

To renal pelvis Loop of Henle

FIGURE 48-10 Animated General regions of filtration, reabsorption, and secretion

glomerular capillaries is at much higher pressure, so more plasma is filtered in the kidney. Several factors contribute to filtration. First, the hydrostatic blood pressure in the glomerular capillaries is higher than in other capillaries. This high pressure is mainly due to the high resistance to outflow presented by the efferent arteriole, which is smaller in diameter than the afferent arteriole (see Fig. 48-9b). A second factor contributing to the large amount of glomerular filtrate is the large surface area for filtration provided by the highly coiled glomerular capillaries. A third factor is the great permeability of the glomerular capillaries. Numerous small pores between the endothelial cells that form their walls make the glomerular capillaries more porous than typical capillaries. The wall of Bowman’s capsule in contact with the capillaries consists of specialized epithelial cells called podocytes. These cells have numerous cytoplasmic extensions called foot processes that cover most of the surfaces of the glomerular capillaries (FIG. 48-11). Foot processes of adjacent podocytes are separated by narrow gaps called filtration slits. The porous walls of the glomerular capillaries and the filtration slits of the podocytes form a filtration membrane that permits fluid and small solutes dissolved in the plasma (including glucose, amino acids, sodium, potassium, chloride, bicarbonate, other salts, and urea) to pass through and become part of the

filtrate. This filtration membrane holds back blood cells, platelets, and most of the plasma proteins. The total volume of blood passing through the kidneys is about 1200 mL per minute, or about one-fourth of the entire cardiac output. As plasma passes through the glomerulus, it loses more than 10% of its volume to the glomerular filtrate. The normal glomerular filtration rate amounts to about 180 L (about 45 gallons) each 24 hours. This is 4.5 times the amount of fluid in the entire body! Common sense tells us that urine could not be excreted at that rate. Within a few moments, dehydration would become lifethreatening.

Reabsorption is highly selective The threat to homeostasis posed by the vast amounts of fluid filtered by the kidneys is avoided by reabsorption. The renal tubules reabsorb about 99% of the filtrate into the blood, leaving only about 1.5 L to be excreted as urine during a 24-hour period. Reabsorption permits precise regulation of blood chemistry by the kidneys. Wastes, excess salts, and other materials remain in the filtrate and are excreted in the urine, while needed substances such as glucose and amino acids are returned to the blood. Each day the tubules reabsorb more than 178 L of water, 1200 g (2.6 lb) of salt, and about 250 g (0.5 lb) of glucose. Most of this, of course, is reabsorbed many times over. Glomerulus Bowman's The simple epithelial cells capsule lining the renal tubule are well Afferent adapted for reabsorbing materials. arteriole Blood cells Their abundant microvilli increase restricted fr om passing through the surface area for reabsorption. These epithelial cells contain nuEndothelial cell of capillary merous mitochondria that provide the energy for running the Red Capillary blood cell pumps that actively transport pores cell materials. Efferent Most (about 65%) of the arteriole Nucleus filtrate is reabsorbed as it passes through the proximal convoluted tubule. Glucose, amino acids, vitamins, and other substances of Podocyte nutritional value are entirely reabsorbed there. Many ions, includFiltration ing sodium, chloride, bicarbonate, slits and potassium, are partially reabsorbed. Some of these ions are actively transported; others are reabsorbed by diffusion. Reabsorption continues as the filtrate passes through the loop of Henle and the distal convoluted tubule. Foot processes Then the filtrate is further concentrated as it passes through the collecting duct that leads to the FIGURE 48-11 Filtration membrane of the kidney renal pelvis. The porous walls of the glomerular capillaries and the filtration slits between podocytes form a filtration Substances that are useful membrane that is highly permeable to water and small molecules but restricts the passage of blood cells and large molecules. to the body, such as glucose or

amino acids, are normally reabsorbed from the renal tubules. If the concentration of a particular substance in the blood is high, however, the tubules may not be able to reabsorb it all. The maximum rate at which a substance can be reabsorbed is its tubular transport maximum (Tm). When that rate is reached, the binding sites are saturated on the membrane proteins that transport the substance. For example, the tubular load of glucose is about 125 mg per minute, and almost all of it is normally reabsorbed. However, in a person with uncontrolled diabetes mellitus, the concentration of glucose in the blood exceeds its Tm (glucose concentration in excess of 320 mg per minute). The excess glucose cannot be reabsorbed and is excreted in the urine (glucosuria), a symptom of diabetes (discussed in Chapter 49). Some substances are actively secreted from the blood into the filtrate Tubular secretion is the selective transfer of substances from the blood in the peritubular capillaries into the renal tubule. Note that in tubular secretion ions and other substances are moved across the tubule epithelium in the opposite direction of reabsorption. Potassium, hydrogen ions, and ammonium ions, as well as some organic ions such as creatinine (a metabolic waste), are secreted into the filtrate. Certain drugs, such as penicillin, are also removed from the blood by secretion. Secretion occurs mainly in the region of the distal convoluted tubule. Secretion of hydrogen ions by the collecting ducts is an important homeostatic mechanism for regulating the pH of the blood. Carbon dioxide, which diffuses from the blood into the cells of the distal tubules and collecting ducts, combines with water, which produces carbonic acid. This acid then dissociates to form hydrogen ions and bicarbonate ions. When the blood becomes too acidic, the collecting ducts secrete more hydrogen ions into the urine: CO2 + H2O Δ H2CO3 Δ H+ + HCO3−

Potassium ion secretion is another important homeostatic mechanism. When K+ concentration is too high, nerve impulses are not effectively transmitted, and the strength of muscle contraction decreases. The heart rhythm becomes irregular, and cardiac arrest can occur. When K+ concentration exceeds its homeostatic level, K+ are secreted from the blood into the renal tubules and then excreted in the urine. Secretion results partly from a direct effect of the K+ on the tubules. In addition, the adrenal cortex (a region of the adrenal gland) increases its output of the hormone aldosterone, which further stimulates secretion of K+.

Urine becomes concentrated as it passes through the renal tubule We can survive with limited fluid intake because the kidneys can produce highly concentrated urine—more than four times as concentrated as blood. The osmolarity of human blood is about 300 milliosmoles per liter (mOsm/L). The kidneys can produce urine with an osmolarity of about 1200 mOsm/L. As the filtrate passes through various regions of the renal tubule, salt (NaCl) is reabsorbed into the interstitial fluid, and a salt

concentration gradient is established (FIGS. 48-12 and 48-13). The gradient is used to produce concentrated urine. When filtrate flows from Bowman’s capsule to the proximal tubule, its osmolarity is the same as that of blood (about 300 mOsm/L). Water and salt are reabsorbed from the proximal tubule. Sodium ions are actively transported out of the proximal tubule, and chloride follows passively. As salt passes into the interstitial fluid, water follows osmotically. The walls of the descending limb of the loop of Henle are relatively permeable to water but relatively impermeable to sodium and urea. The interstitial fluid has a high concentration of Na+, so as the filtrate passes down the loop of Henle, water moves out by osmosis. This process concentrates the filtrate inside the loop of Henle. The loop of Henle is specialized to highly concentrate sodium chloride in the interstitial fluid of the medulla. It maintains a highly hypertonic interstitial fluid in the medulla near the bottom of the loop, which in turn lets the kidneys produce concentrated urine. At the turn of the loop of Henle, the walls become more permeable to salt and less permeable to water. As the concentrated filtrate begins to move up the ascending limb (the thin region), salt diffuses out into the interstitial fluid. This contributes to the high salt concentration of the interstitial fluid in the medulla surrounding the loop of Henle. Farther along the ascending limb (the thick region), sodium is actively transported out of the tubule. Because water passes out of the descending limb of the loop of Henle, the filtrate at the bottom of the loop has a high salt concentration. However, because salt (but not water) is removed in the ascending limb, by the time the filtrate moves into the distal tubule, its osmolarity may be the same as or even lower than that of blood. As it passes through the distal tubule, the filtrate may become even more dilute. The distal tubule is relatively impermeable to water but actively transports salt out into the interstitial fluid. The filtrate passes from the renal tubule into a larger collecting duct that eventually empties into the renal pelvis. Note the counterflow of fluid through the two limbs of the loop of Henle. Filtrate passing down through the descending limb is flowing in a direction opposite that of the filtrate moving upward through the ascending limb. The filtrate becomes concentrated as it moves down the descending limb and diluted as it moves up the ascending limb. This countercurrent mechanism helps maintain a high salt concentration in the interstitial fluid of the medulla. The hypertonic interstitial fluid draws water osmotically from the filtrate in the collecting ducts. The inner medullary collecting ducts are permeable to urea, so some of the concentrated urea in the filtrate can diffuse out into the interstitial fluid. The urea contributes to the high solute concentration of the inner medulla. This process helps concentrate urine. The collecting ducts pass through the zone of very salty interstitial fluid. As the filtrate moves through the collecting duct, water passes osmotically into the interstitial fluid. Excess interstitial fluid continuously enters capillaries and becomes part of the blood. Sufficient water can leave the collecting ducts to produce a highly concentrated urine. A hypertonic urine conserves water. Some of the water that diffuses from the filtrate into the interstitial fluid is removed by the vasa recta, which are long,

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FIGURE 48-12 Animated Movement of water, ions, and urea through the renal tubule and collecting duct Water passes out of the descending limb of the loop of Henle, leaving a more concentrated filtrate inside. The heavy outline along the ascending limb indicates that this region is relatively impermeable to water. NaCl diffuses out from the lower (thin) part of the ascending limb. In the upper (thick) part of the ascending limb, NaCl is actively transported into the interstitial fluid. The saltier the interstitial fluid becomes, the more water moves out of the descending limb. This process leaves a concentrated filtrate inside, so more salt passes out. Note that this is a positive feedback system. Some urea also moves out into the interstitial fluid through the collecting ducts. Water from the collecting ducts moves out osmotically into this hypertonic interstitial fluid and is carried away by capillaries.

Afferent arteriole

Bowman's capsule

Efferent arteriole

Proximal tubule

Distal tubule

NaCl

H2O

H2O

Filtrate

CORTEX MEDULLA

NaCl NaCl NaCl

H2O

Collecting duct

H2O H2O

NaCl Urea

Descending limb

Ascending limb Loop of Henle

straight capillaries that extend from the efferent arterioles of the juxtamedullary nephrons. The vasa recta extend deep into the medulla, only to negotiate a hairpin curve and return to the cortical venous drainage of the kidney. These capillaries parallel the renal tubul